Skip to main content

Full text of "Revue suisse de zoologie"

See other formats


Uhanbidgs aris 
stiva 


2424372 
ine 


Eee 
terrors 


zu: 
teo: 
ee 
Als. binc4.0.4/ 
Alea: 


IPRFEPETLOLI 
PESI a4 ia) 


Se hae, 
Soe R ay 
i iù 


Uae GLA, 


Lat eek 


Pr PT 


OCT 


PATATE 


NPA 
Venen, u 
iyafentein 4 
ren PETE LIRE BI 
RTE 
nyt tak Cote 


en 

Sisti 

seni sine 

u TSI COLO 
nt 


i Ä 
HET 
Rte 
Mi 4 erty x KERLE N 
VRR re I TELL i CRIME 
7 a 


REVUE SUISSE 


DE 


ZOOLOGIE 


REVUE SUISSE 
ZOOLOGIE 


ANNALES 


DE LA 
SOCIETE SUISSE DE ZOOLOGIE 
ET DU 


MUSEUM D'HISTOIRE NATURELLE 
DE LA VILLE DE GENEVE 


la 


GENEVE 
2000 


ISSN 0035-418X 


TABLE DES MATIERES 


TOME 107 — 2000 


Fascicule 1 


JUVARA-BALS, Ilinca & Vojciech WITALINSKI. Description of five new species of 
Holoparasitus s. str. with redescription of H. apenninorum (Berlese, 1906) 
and H. cultriger (Berlese, 1906) from Italy and Spain (Acari, Gamasida, 
Barasıldao) Re AN RI DEA N ikl ee LR URI 


SILVA, Carla Maria Menegola da & Beatriz MOTHES. Three new species of Geodia 
Lamarck, 1815 (Porifera, Demospongiae) from the bathyal depths off 
Brazilianicoast.s outhwestem- AtlantiGe EER a een 


MAHUNKA, Sandor & Luise MAHUNKA-PAPP. Oribatids from Switzerland III 
(Acari: Oribatida: Oppiidae | and Quadroppiidae). (Acarologica Genavensia 


GIL DE PERTIERRA, Alicia A. & Alain de CHAMBRIER. Rudolphiella szidati sp. n. 
(Proteocephalidea: Monticelliidae, Rudolphiellinae) parasite of Luciopi- 
melodus pati (Valenciennes, 1840) (Pisces: Pimelodidae) from Argentina 
with new observations on Rudolphiella lobosa (Riggenbach, 1895)........ 


BENJAMIN, Suresh P. & Rudy JOCQUÉ. Two new species of the genus Suffasia 
Pomorie wanka (Araneae: ZOd arid ae) se een me et ee ee en ee) stat IE 


LANG, Claude. Diversité du zoobenthos dans 47 rivières du canton de Vaud: 
tendances ES 1 O ieee ee VOIR En 


Vit, Stanislav. Contribution a la connaissance de la famille Eucinetidae (Cole- 
Oi OUST) ER N SR E PE Er IE a a RR NEE 


BANZIGER, Ruth. Spatio-temporal distribution of size classes and larval instars of 
aquatic insects (Ephemeroptera, Trichoptera and Leopidoptera) in a Potamo- 
ecton pecinatus ds bed (Make'Geneva, Switzerland)... 2.20... RENE 


BESUCHET, Claude & Stanislav Vit. Les Nanophthalmus Motschulsky d’Europe 
(@oleoptera,seydmaenidae) ZERI 


Heyp, Andreas & Wolfgang PFEIFFER. Uber die Lauterzeugung der Welse (Silu- 
roidei, Ostariophysi, Teleostei) und ihren Zusammenhang mit der Phylo- 
sentetundsderSchreekreaktions rn A ey eae 


GANTENBEIN, Benjamin, Christian KRoPF, Carlo Rodolfo LARGIADER & Adolf 
SCHOLL. Molecular and morphological evidence for the presence of a new 
Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus............. 


Fascicule 2 


LANG, Claude. Etat trophique du lac de Morat indiqué par le zoobenthos: tendance 
LO SOB OS eae Maia arenes as nr NEST ae Se nO 


AZPELICUETA, Maria de la Mercedes & José O. GARCIA. A new species of Astya- 
nax (Characiformes, Characidae) from Uruguay river basin in Argentina, 
withrremarkssonthooksptesenceiin@haracıdaege nn. m ar 


Pages 


3-30 


31-48 


49-79 


81-95 


97-106 


107-122 


123-138 


139-151 


153-163 


165-211 


213-232 


233-243 


245-257 


VI TABLE DES MATIERES 


BALLERIO, Alberto. Revision of the genus Ebbrittoniella Martinez (Coleoptera: 
Scarabacoideay Ceratocanthidae) 4... RE 


LIENHARD, Charles. A new desert psocid from Namibia (Insecta: Psocoptera: 
NiO SUN AS) Bene ee ise ists dle: ee ee eee ee 


LEISTIKOW, Andreas. Terrestrial Isopoda from Guatemala and Mexico (Crustacea: 
OniscideaxErimocheta eye se eee eee Cee eee 


BURCKHARDT, Daniel & Mariängela GUAJARA. Euphalerus clitoriae sp. n., a new 
psyllid species from Clitoria fairchildiana (Fabaceae, Papilionoideae), and 
notes on other Euphalerus spp. (Hemiptera, Psylloidea)................. 


Koss, Kerstin & Wolfgang PFEIFFER. Zur Biologie des “Eigenweidefisches” 
Carapus acus (Briinnich, 1768) (Carapidae, Teleostei), mit Hinweisen auf 
einennicht-parasitische’Ernährung® PEER RR eee 


BEDOS, Anne & Louis DEHARVENG. Un nouveau Collembole Neanurinae du Sud 


du Vietnam, Blasconura batai sp. n., avec une clé des especes du genre 
(@ollembolaxNeanurrdao)e NS ee E TORE DA o REA 


BAUR, Hannes & Felix AMIET. Die Leucospidae (Hymenoptera: Chalcidoidea) der 
Schweiz, mit einem Bestimmungsschlüssel und Daten zu den europäischen 


TANDY, Mills & Jean-Luc PERRET. The Bufo tuberosus species group with the 
description of a new species from the rainforest of Cöte-d’Ivoire.......... 


Fascicule 3 


PUTHZ, Volker. The genus Dianous Leach in China (Coleoptera, Staphylinidae). 
2OIAEontibutionitoitheiknowledse.of Steninae. 222. a. er sar ee 


GATTOLLIAT, J.-L. & M. SARTORI. Contribution to the systematics of the genus 
Dabulamanzia (Ephemeroptera: Baetidae) in Madagascar. ............... 


MUSTER, Christoph & Konrad THALER. Das Männchen von Zelotes zellensis 
Gurmmı(Araneae:/Gnaphosidae) er. LELE 


VOGEL, Peter, Mark LAWRENCE & Ali AGHNAJ. Note sur les musaraignes 
(Soricidae, Mammalia) du Pare National du Souss-Massa, Maroc.......... 
Lost, Ivan. A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the 
ReoplesssRepubliciof@hina; Mer IT aoe eee 


Fascicule 4 


POMORSKI, Romuald J. & Dariusz SKARZYNSKI. Redescription of Hymenaphorura 
alticola (Bagnall, 1935) from the Alps and description of a new related 
species from the Sudetes, Hymenaphorura improvisa sp. n. (Collembola: 
OnychiunGae) i. lie Sa su. be oe ss dug +s) ae eee 


ZAWADZKI, Claudio H., Roberto E. Reis & Erasmo RENESTO. Allozyme discri- 
mination of three species of Loricariichthys (Siluriformes: Loricariidae) from 
SouMenM Brazile dg. (000 NORTE 


Pages 


259-275 


277-281 


283-323 


325-334 


335-349 


351-357 


359-388 


389-418 


419-559 


561-577 


579-589 


591-599 


601-656 


657-662 


663-674 


TABLE DES MATIERES 


MAHUNKA, Sandor. Oribatids from Sabah (East Malaysia) VIII (Acari: Oribatida: 
Dampfiellidae and Otocepheidae). (Acarologica Genavensia LXXXVI)...... 


LOURENÇO, Wilson R. More about the Buthoidea of Madagascar, with special 
references to the genus Tiryobuthus Pocock (Scorpiones, Buthidae)......... 


PAGES, Jean. Japygidés (Diplura) du Sud-Est asiatique n° 9. - Dicellurata Gena- 
DENSO PROIETTI REA EA a ASE ER Ne Le 


COMELLINI, André. Notes sur les Psélaphines néotropicaux (Coleoptera, Staphy- 
linidae, Pselaphinae) — 11. Un nouveau genre et sept espèces nouvelles de la 
GRIDUEGESYMIELOPIASIM PE SE an PAUL ee ee 


BÖHME, Wolfgang, Andreas SCHMITZ & Thomas ZIEGLER. A review of the West 
African skink genus Cophoscincopus Mertens (Reptilia: Scincidae: Lygoso- 
minae): resurrection of C. simulans (Vaillant, 1884) and description of a new 
SPECIES IE LE ee N ee 


ROWELL, C. H. F. Review of the Lithoscirtus genus group (Orthoptera, Acrididae, 
Rroctolabmaec)) with description of mew species: ... 2.0... 00 ee ve eee 


VALLAN, Denis. A new species of the genus Stumpffia (Amphibia: Anura: 
Microhylidae) from a small forest remnant of the central high plateau of 
Madagascar Sewerage cit Reese le scoot tua RN ce co ER 


GANTENBEIN, Benjamin, Victor FET, Mark BARKER & Adolf SCHOLL. Nuclear and 
mitochondrial markers reveal the existence of two parapatric scorpion species 
in the Alps: Euscorpius germanus (C. L. Koch, 1837) and E. alpha 
Caponaccoy 195 04statenov. (Euscorpiidac)ia- ea ale 


LIENHARD, Charles. A new genus of Prionoglarididae from a Namibian cave 
(insectasPSOCoptera) se ar. NE EN <a NE WB REN eo RE 


MERZ, Bernhard. Two new Tephritidae (Diptera) from the Western Palaearctic 
ICROM les SE cet SRR TOI OT NE ate EEE ER N ar 


VII 


Pages 


675-720 


721-736 


737-764 


765-776 


777-791 


793-834 


835-841 


843-869 


871-882 


883-892 


DR 
N 
Lu 


INDEX DES AUTEURS 
par 


ORDRE ALPHABETIQUE 


AZPELICUETA, Maria de la Mercedes & José O. GARCIA. A new species of Astya- 
nax (Characiformes, Characidae) from Uruguay river basin in Argentina, 
with remarks on hook presence in Characidae. ........................ 


BALLERIO, Alberto. Revision of the genus Ebbrittoniella Martinez (Coleoptera: 
Scarabacoideas Cerato Cani dI) EROI ee: 


BANZIGER, Ruth. Spatio-temporal distribution of size classes and larval instars of 
aquatic insects (Ephemeroptera, Trichoptera and Leopidoptera) in a Potamo- 
geton pectinatus L. bed (Lake Geneva, Switzerland). ................... 


BAUR, Hannes & Felix AMIET. Die Leucospidae (Hymenoptera: Chalcidoidea) der 
Schweiz, mit einem Bestimmungsschliissel und Daten zu den europäischen 


BEDOS, Anne & Louis DEHARVENG. Un nouveau Collembole Neanurinae du Sud 
du Vietnam, Blasconura batai sp. n., avec une clé des especes du genre 
(GollembolasiNeanunid ae) yes ee oes PR eine 


BENJAMIN, Suresh P. & Rudy JOCQUÉ. Two new species of the genus Suffasia 
fromesnilcankal(Aranecae=Zodarlidae) I re alle eect tie INDI one 


‘ BESUCHET, Claude & Stanislav VIT. Les Nanophthalmus Motschulsky d’Europe 
(ColeopteratScydmacnidae)y et. acess ese n 


BÖHME, Wolfgang, Andreas SCHMITZ & Thomas ZIEGLER. A review of the West 
African skink genus Cophoscincopus Mertens (Reptilia: Scincidae: Lygoso- 
minae): resurrection of C. simulans (Vaillant, 1884) and description of a new 
SPECIES See a STIRIA a Re ap OC AE MNT Al Ra AE 


BURCKHARDT, Daniel & Mariangela GUAJARA. Euphalerus clitoriae sp. n., a new 
psyllid species from Clitoria fairchildiana (Fabaceae, Papilionoideae), and 
notes on other Euphalerus spp. (Hemiptera, Psylloidea). ................ 


COMELLINI, André. Notes sur les Psélaphines néotropicaux (Coleoptera, Staphy- 
linidae, Pselaphinae) — 11. Un nouveau genre et sept espèces nouvelles de la 
teibusdessMetopiasını 1.05 ar. EE IO TI 


GANTENBEIN, Benjamin, Christian Kropr, Carlo Rodolfo LARGIADER & Adolf 
SCHOLL. Molecular and morphological evidence for the presence of a new 
Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus............. 


GANTENBEIN, Benjamin, Victor Fer, Mark BARKER & Adolf SCHOLL. Nuclear and 
mitochondrial markers reveal the existence of two parapatric scorpion species 
in the Alps: Euscorpius germanus (C. L. Koch, 1837) and E. alpha 
Caponiaccoml 950 statenovec(2usconpidae) a) er II ER 


GATTOLLIAT, J.-L. & M. SARTORI. Contribution to the systematics of the genus 
Dabulamanzia (Ephemeroptera: Baetidae) in Madagascar............... 


245-257 


259-275 


139-151 


359-388 


351-357 


97-106 


153-163 


777-791 


325-334 


765-776 


843-869 


561-577 


x INDEX DES AUTEURS 


GIL DE PERTIERRA, Alicia A. & Alain de CHAMBRIER. Rudolphiella szidati sp. n. 
(Proteocephalidea: Monticelliidae, Rudolphiellinae) parasite of Luciopi- 
melodus pati (Valenciennes, 1840) (Pisces: Pimelodidae) from Argentina 
with new observations on Rudolphiella lobosa (Riggenbach, 1895)........ 


Heyp, Andreas & Wolfgang PFEIFFER. Über die Lauterzeugung der Welse (Silu- 
roidei, Ostariophysi, Teleostei) und ihren Zusammenhang mit der Phylo- 
genieund: der Schreckreaktion. sto. IRE OS PERE 


JUVARA-BALS, Ilinca & Vojciech WITALINSKI. Description of five new species of 
Holoparasitus s. str. with redescription of H. apenninorum (Berlese, 1906) 
and H. cultriger (Berlese, 1906) from Italy and Spain (Acari, Gamasida, 
Barasiudao)e E LORIA RI 


KLoss, Kerstin & Wolfgang PFEIFFER. Zur Biologie des “Eigenweidefisches” 
Carapus acus (Briinnich, 1768) (Carapidae, Teleostei), mit Hinweisen auf 
einemicht-parasitische Ernährung... LO RE 


LANG, Claude. Diversité du zoobenthos dans 47 rivières du canton de Vaud: 
tendaneesl 98 NIIT, Ve ee A EE RS 


LANG, Claude. Etat trophique du lac de Morat indiqué par le zoobenthos: tendance 
OS FOO STE An RE i MR en ee 


LEISTIKOW, Andreas. Terrestrial Isopoda from Guatemala and Mexico (Crustacea: 
OniscideasGninocheta) a. Sr sae oe ET oe RI 


LIENHARD, Charles. A new desert psocid from Namibia (Insecta: Psocoptera: 
Miro sida) arg... sini We Seen ee news a, + CORE 


LIENHARD, Charles. A new genus of Prionoglarididae from a Namibian cave 
(InsectaaBsocoptera) at ila e die CES RE 


LößL, Ivan. A review of the Scaphidiinae (Coleoptera: Staphylinidae) of the 
ReoplessiRepubliciof China IE RER ERI RR 


LOURENÇO, Wilson R. More about the Buthoidea of Madagascar, with special 
references to the genus Tiryobuthus Pocock (Scorpiones, Buthidae)......... 
MAHUNKA, Sandor & Luise MAHUNKA-PAPP. Oribatids from Switzerland III 
(Acari: Oribatida: Oppiidae | and Quadroppiidae). (Acarologica Genavensia 


MAHUNKA, Sandor. Oribatids from Sabah (East Malaysia) VIII (Acari: Oribatida: 
Dampfiellidae and Otocepheidae). (Acarologica Genavensia LXXXVI)...... 


MERZ, Bernhard. Two new Tephritidae (Diptera) from the Western Palaearctic 
RAT BE ne Cu à eee ne LR PE PNR Mla cin 5.0.0056 0° 


MUSTER, Christoph & Konrad THALER. Das Männchen von Zelotes zellensis 
Grimm(Araneae:/Gnaphosidae)!.... 2... NOR fae oe ee eee 
PAGES, Jean. Japygidés (Diplura) du Sud-Est asiatique n° 9. - Dicellurata Gena- 
VERSO PON IN nenne release IE 
POMORSKI, Romuald J. & Dariusz SKARZYNSKI. Redescription of Hymenaphorura 
alticola (Bagnall, 1935) from the Alps and description of a new related 
species from the Sudetes, Hymenaphorura improvisa sp. n. (Collembola: 
Ouychiundae): rn Ni os ches Se ee SE EEE 
PUTHZ, Volker. The genus Dianous Leach in China (Coleoptera, Staphylinidae). 
2618 Contribution! to the knowledge of Steninaer .. 2.22. mer een 


ROWELL, C. H. F. Review of the Lithoscirtus genus group (Orthoptera, Acrididae, 
Proctolabinae)) with description of new) Species; Re CPP EC EE 


Pages 


81-95 


165-211 


3-30 


335-349 


107-122 


233-243 


283-323 


277-281 


871-882 


601-656 


721-736 


49-79 


675-720 


883-892 


579-589 


737-764 


657-662 


419-559 


793-834 


INDEX DES AUTEURS 


SILVA, Carla Maria Menegola da & Beatriz MOTHES. Three new species of Geodia 
Lamarck, 1815 (Porifera, Demospongiae) from the bathyal depths off 
Braziliankcoase SouthwestemirAtlantt Cae ee Zur OO 


TANDY, Mills & Jean-Luc PERRET. The Bufo tuberosus species group with the 
description of a new species from the rainforest of Cote-d’Ivoire.......... 


VALLAN, Denis. A new species of the genus Stumpffia (Amphibia: Anura: 
Microhylidae) from a small forest remnant of the central high plateau of 
Madagascar EA ee na te eer RAO wie RENE 

VIT, Stanislav. Contribution à la connaissance de la famille Eucinetidae (Cole- 
UE TAN) Nee Paste ERNEST meee et SEAN Mi aL EM ete a eue ve EN ON 

VOGEL, Peter, Mark LAWRENCE & Ali AGHNAJ. Note sur les musaraignes 
(Soricidae, Mammalia) du Parc National du Souss-Massa, Maroc......... 

ZAWADZKI, Claudio H., Roberto E. Reis & Erasmo RENESTO. Allozyme discri- 
mination of three species of Loricariichthys (Siluriformes: Loricariidae) from 
Southe mp raza se ee Nest 


XI 


31-48 


389-418 


835-841 


123-138 


591-599 


663-674 


ICIETE SUISSE DE ZOOLOGIE 
ÉUM D'HISTOIRE NATURELLE 


AGMHSONIA 


| RS 2000 ISSN 0035 - 418 X 


SWISS JOURNAL OF ZOOLOGY 


REVUE SUISSE DE ZOOLOGIE 


REVUE SUISSE DE ZOOLOGIE 


TOME 107 — FASCICULE | 


Publication subventionnée par: 


ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN 
VILLE DE GENEVE 
SOCIETE SUISSE DE ZOOLOGIE 


VOLKER MAHNERT 
Directeur du Muséum d'histoire naturelle de Genève 


MANUEL RUEDI 
Chargé de recherche au Muséum d’histoire naturelle de Genève 


CHARLES LIENHARD 
Chargé de recherche au Muséum d'histoire naturelle de Geneve 


Comité de lecture 


Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des Instituts de zoologie des universités 


suisses. 


Les manuscrits sont soumis à des experts d’ institutions suisses ou étrangères selon le 


sujet étudié. 


La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 


comparée, physiologie. 


Administration 
MUSEUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 


Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 


PRIX DE L'ABONNEMENT: 


(en francs suisses) 


Les demandes d'abonnement doivent être adressées 
à la rédaction de la Revue suisse de Zoologie, 
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 


SUISSE Fr. 225.— UNION POSTALE Fr. 230.— 


ANNALES 


de la 

SOCIETE SUISSE DE ZOOLOGIE 

et du 

MUSEUM D'HISTOIRE NATURELLE 
de la Ville de Geneve 


tome 107 
fascicule 1 
2000 


E à 
kl GENEVE MARS 2000 ISSN 0035 - 418 X 


SWISS JOURNAL OF ZOOLOGY 


REVUE SUISSE DE ZOOLOGIE 


REVUE SUISSE DE ZOOLOGIE 


TOME 107 — FASCICULE 1 


Publication subventionnée par: 
ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN 
VILLE DE GENEVE 
SOCIETE SUISSE DE ZOOLOGIE 


VOLKER MAHNERT 
Directeur du Muséum d'histoire naturelle de Genève 


MANUEL RUEDI 
Charge de recherche au Muséum d’histoire naturelle de Geneve 


CHARLES LIENHARD 
Chargé de recherche au Muséum d'histoire naturelle de Genève 


Comité de lecture 


Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Geneve et de représentants des Instituts de zoologie des universités 
suisses. 

Les manuscrits sont soumis a des experts d’institutions suisses ou étrangeres selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 
comparée, physiologie. 


Administration 


MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 


Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 


PRIX DE L'ABONNEMENT: 


SUISSE Fr. 225.— UNION POSTALE Fr. 230.— 


| (en francs suisses) 


Les demandes d'abonnement doivent étre adressées 
à la rédaction de la Revue suisse de Zoologie, 
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 


REVUE SUISSE DE ZOOLOGIE 107 (1): 3-30; mars 2000 


Description of five new species of Holoparasitus s. str. with 
redescription of H. apenninorum (Berlese, 1906) and A. cultriger 
(Berlese, 1906) from Italy and Spain (Acari, Gamasida, Parasitidae) 


Ilinca JUVARA-BALS! & Wojciech WITALINSKI? 

| Museum of Natural History, CP 6434, CH-1211 Geneva 6, Switzerland. 

? Institute of Zoology, Jagiellonian University, Ingardena 6, PL-30060 Krakow, 
Poland. 


Description of five new species of Holoparasitus s. str. with redes- 
cription of H. apenninorum (Berlese, 1906) and H. cultriger (Berlese, 
1906) from Italy and Spain (Acari, Gamasida, Parasitidae). - The spe- 
cimens included under the name of H. apenninorum (Berlese, 1906) in the 
Berlese collection in Florence are reviewed and a lectotype for H. apen- 
ninorum is designated. Five new species, H. cornutus sp. n., H. crassi- 
setosus Sp. n., H. digitiformis sp. n., A. ellipticus Sp. n., H. gibber sp. n. are 
described and H. cultriger (Berlese, 1906) is redescribed, all from material 
from Italy and Spain. The characteristics of a new species group mallorcae 
is given. 


Key-words: Acari - Gamasida - Parasitidae - Holoparasitus - taxonomy. 


INTRODUCTION 


The gamasids belonging to the genus Holoparasitus Oudemans, 1936 are free 
living, ground inhabiting predatory mites, distributed in the Holarctic region. This 
genus comprises 34 species which are divided, accordingly to Juvara-Bals (1975) and 
Hyatt (1987), into three subgenera: Holoparasitus s. str. (30 species), Heteroparasitus 
Juvara-Bals, 1975 (3 species) and Ologamasiphis Holzmann, 1969 (1 species). 

In the subgenus Holoparasitus s. str., Micherdzifiski (1969) distinguished two 
species-groups: calcaratus and pollicipatus-groups, whereas the remaining species are 
considered as incertae sedis. Juvara-Bals (1975) proposed another group of species, 
named caesus-group. More recently Hyatt (1987) mentioned only the species-groups 
recognized by Micherdzinski and designated a neotype for H. calcaratus (C.L. Koch, 
1839), the type species of genus Holoparasitus. 

During the last decade new taxa have been identified in the course of faunistic 
and ecological programs or from acarological collections (Schmölzer, 1991, 1995a, 
1995b; Vinnik, 1994; Witalinski, 1994a, 1994b: Juvara-Bals, 1995). As mentioned 
above, Holoparasitus s. str. includes now 30 species. Unfortunately, some of these 


Manuscript accepted 27.10.1999 


4 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


species are known incompletely (one sex only) and their descriptions frequently lack 
details. This rather large subgenus strongly requires a phylogenetical analysis. 
However, it cannot be successfully done until the following problems are still unsolved: 


- the revision of the species deposited in the collections of the Berlese, v. 
Vitzthum and Willmann; 

~ the re-examination of some key morphological characters omitted in early 
descriptions in some species; 

~ a more complete documentation about the gamasid mites in many European 
regions, especially those from south-east and central Europe, is available. 


The aim of this paper is to advance the knowledge of species included in the 
subgenus Holoparasitus s. str. as defined in Juvara-Bals (1975). A revision of spe- 
cimens labelled as A. apenninorum (Berlese, 1906) in the Berlese collection (“Berlese 
Acaroteca”, Florence, Italy) showed that this is a complex of four species: one of 
them we have identified as the nominal species while the three others are new species. 
We also describe two additional new species from Spain and Sicily, H. gibber sp. n. 
and A. ellipticus sp. n., respectively. These latter species form together with A. sicu- 
lus (Berlese, 1905), H. mallorcae Juvara-Bals, 1975, H. lawrencei Hyatt, 1987 and 
H. maritimus Hyatt, 1987, a well defined group of species which we named mallor- 
cae-group. Additionally, we redescribe the male of A. cultriger (Berlese, 1906) based 
on the single type specimen in the Berlese Acaroteca. 


MATERIAL AND METHODS 


The material comes from Italy and Spain. One of us (WW) was provided with a 
large collection of samples from Italy collected by Prof. R. Dallai and the staff of the 
Department of Evolutionary Biology, University of Siena, Italy (DEBS). Juvara-Bals 
had the opportunity to identify samples of mites (family Parasitidae) from Italy 
(Toscana) collected by Dr. F. Pegazanno and Dr. R. Nannelli from the Experimental 
Institute of Agricultural Zoology (EIAZ), Florence, Italy. She was also working on the 
genus Holoparasitus from the Athias-Henriot collection deposited in the Museum of 
Natural History (MHNG), Geneva, Switzerland. 

We studied material from the Berlese Acaroteca (EIAZ) which contains several 
species collected generally around Florence (Toscana) and in Umbria. 

Morphological terminology follows mainly Evans and Till (1979) and Van der 
Hammen (1980). Setal notation for the idiosoma follows Lindquist and Evans (1965). 
Measurements were made from slide-mounted material and expressed in micrometers 
(um). The types are deposited in the Zoological Museum of the Jagiellonian University, 
Cracow, Poland (ZMJU), in the MHNG and in the EIAZ. 


Fic. | 


Holoparasitus apenninorum (Berlese,1906). Male: A- chelicera, paraxial view; B-idem, antiaxial: 
C-pedipalp, trochanter and femur; D-corniculus; E-leg II, femur, genu and tibia. 

H. cultriger (Berlese,1906). Male: F-corniculus; G-chelicera, paraxial; H-sternogenital region and 
genital lamina; I-leg II, femur, genu and tibia; J-tectum (after Berlese, 1906). 


NEW SPECIES OF HOLOPARASITUS 5 


6 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


SYSTEMATIC ACCOUNT 


REVISION OF HOLOPARASITUS APENNINORUM (BERLESE, 1906) AND H. CULTRIGER 
(BERLESE, 1906) 


The material of H. apenninorum (Berlese, 1906) in the Berlese collection 
named Ologamasus pollicipatus var. apenninorum consists of 28 specimens on 10 
slides and is in fact a mixture of four species, 1.e.: 


1. H. apenninorum (Berlese, 1906), 16 non-dissected (slide 5/19), Vallom- 
brosa (Toscana, Italy), from moss; lectotype by present designation. 

2. H. crassisetosus sp. n., 19 dissected (6/40), 49 2 (6/41), 688, 1% 
2 deutonymphs (7/35), 26 4, 29 9 (7/37), Vallombrosa. 

3. H. digitiformis sp. n., 12 (6/42), 15 (6/43), 12 (6/44), Monte Senario 
(Toscana). 

4. H. cornutus sp. n., 16, 12 (7/39), Bevagna (Umbria, Italy); 16, 39 2 
(7/40), Monte alle Forche (Toscana). 

Berlese also recognized in his material from Vallombrosa another variety, 
Ologamasus pollicipatus var. cultriger. This is a valid species, Holoparasitus cultri- 
ger (Berlese,1906), though only one specimen is known, a dissected male which is on 
slide 5/19 together with the lectotype of H. apenninorum noted above. Some speci- 
mens of Paragamasus decipiens (Berlese, 1903) and of Holoparasitus mentioned 
above are mounted together: 1 2 on slide 6/44, 1 2 on slide 7/39 and 16, 19 on slide 
7/40. 


H. apenninorum (Berlese, 1906) Fig. 1 A-E 


Gamasus (Ologamasus) pollicipatus var. apenninorum Berlese, 1906: 253 (in part). 
Holoparasitus lichenis var. apenninorum (Berlese, 1887) sensu Turk 1953, Micherdzifiski 
196972; 

not Holoparasitus apenninorum (Berlese, 1906) sensu Juvara-Bals 1975, Acarologia, p: 400. 


Type material: lectotype 1d non-dissected, slide 5/19, from moss, Vallombrosa (Tos- 
cana) in Berlese Acaroteca, EIAZ; by present designation. 

Our description is based on a non-dissected single male in lateral position so 
that only some of the main characters could be observed. 

Diagnosis. Male: excipulum absent; cheliceral movable digit with a single tooth, 
fixed digit enlarged along its distal third, with 6-7 denticles; leg II with the femoral 
apophysis straight and thumb-like (Fig. IE ). 

Description. Male. The legs I-IV were attached only on the right side of the 
animal. The others were lost but leg II could be seen in the mounting medium detached 
from the animal. Gnathosoma. Movable digit of chelicera with a single tooth situated 
medially. Fixed digit, enlarged along its upper third, with 6-7 denticles. Spermatotreme 
in the form of a fine slit, arthrodial process setiform antiaxially and brush-like 
paraxially (Fig. | A,B). Pedipalp: trochanter simple, femur with slight ventral protu- 
berance located distally (Fig. 1 C). Corniculi triangular, with small protuberance on 
ventral side (Fig. | D). 


NEW SPECIES OF HOLOPARASITUS 7 


Leg II. Main features of leg II shown in figure 1 E. Femoral apophysis straight 
and thumb-like, axillary process elongated and smooth. Spur on genu and tibia conical, 
located midway on ventral face. Measurements: tarsus | = 104 um; tarsus IV = 122um. 

Discussion. Berlese’s (1906) original description of H. apenninorum was based 
on a male and several females, from a moss sample taken at Vallombrosa. Berlese 
described in detail only the male’s chelicera but presented figures of three different 
chelicerae as well as of the epistome, and of the male’s leg II. The drawing of the 
chelicera shown in the fig. 15a, tab. XIII. (6 from Vallombrosa) corresponds to the 
original description which specified that “digitus mobilis basi latiusculus, externe 
gibbosus, dente magno, unico...”; the characteristics of this type of chelicera can be 
seen only on the non-dissected male of the slide 5/19. Berlese attributed to H. apenni- 
norum two other chelicerae (Fig. 15, Fig. 16, tav. XIII) which have two teeth on the 
inner margin of the digitus mobilis. He also mentioned a figure 16a which in fact does 
not exist. Micherdzinski (1969) paid also attention to the similarity between Berlese’s 
initial description and only one of the chelicerae illustrated. Berlese’s figure of the 
male’s leg II corresponds either to the leg of H. digitiformis or to that of H. crassiseto- 
sus, two species found in the same area as H. apenninorum. The triangular epistome 
figured by Berlese can not be seen any more because of the lateral position of the 
animal. 

The characteristics of the female endogynium was neither described nor drawn 
in detail; figure 7, tab. XIX refers either to H. digitiformis or to H. crassisetosus which 
have very similar females. Berlese’s material also contained H. crassisetosus from 
Vallombrosa and H. digitiformis from Monte Senario. Unfortunately, we did not find 
another male of H. apenninorum or a corresponding female. However, it is obvious that 
the male mounted on slide 5/19 is a syntype. We designate it as the lectotype of H. 
apenninorum (Berlese, 1906). 

Specimens identified by Juvara-Bals (1975) as H. apenninorum belongs in fact 
to H. cornutus sp. n. The species was misidentified because at that time it was not 
recognized that original description of Berlese related to a mixture of species. 


H. cultriger (Berlese, 1906) Fig. I F-J 


Type material: 16 holotype, dissected (slide 5/19), Vallombrosa (Toscana, Italy), from 
moss, Berlese Acaroteca (EIAZ). 

Diagnosis. Male: excipulum present; cheliceral movable digit with two little 
denticles, fixed digit markedly longer than movable one, blade-like and toothless; leg II 
with tibia bearing two processes: dorsally one rounded located near proximal margin 
and ventally an elongated situated near distal margin (Fig. 1 D. 

Description. Male. The length of the idiosoma could not be measured because 
the single specimen is in pieces. The few characteristics observed are shown in figure 1. 
Sternogenital region reticulated, with slightly sclerotized excipulum; anterior margin of 
genital lamina with several small denticles in the middle (Fig. | H). 

Gnathosoma. Tectum trispinate, central prong well developed (Fig. 1 J). Corni- 
culi triangular, each with ventral protuberance (Fig. 1 F). Chelicera as in figure 1 G: 


8 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


fixed digit blade-like and toothless, markedly longer than movable one; movable digit 
with two little denticles subapically. 

Legs. Armature of legs II as in figure 1 I. Femoral apophysis strong, thumb-like 
and curved; axillary process short and rounded. Genual process large, plump or 
rounded, situated in distal half on ventral face. Tibia with two protuberances: regular 
process elongated and located ventrally close to distal margin of segment, an extra 
process larger, located dorsally on proximal third of segment. Ventral face of trochanter 
IV with protuberance. Measurements: tarsus I = 138 um, tarsus IV = 150 um. 

Discussion. Berlese described this species on the basis of a single male in poor 
condition. The type specimen is together with that of H. apenninorum on the same 
slide. The two types may be easily distinguished: H. apenninorum is in lateral view and 
non-dissected while H. cultriger is in several pieces. The most valuable character 
separating H. cultriger from the other species with an excipulum is the special blade- 
shaped fixed digit of the chelicera. 


Holoparasitus crassisetosus sp. n. Figs. 2,3 


Type material: 13 holotype, 486 d, 669 © paratypes, Vallombrosa (Toscana, Italy) 
(alt. 960 m), coniferous forest (Abies alba), 6.11.1982. Collected by the staff of DEBS. The 
holotype as well as 46 d and 59 © paratypes have been deposited in MHNG, 446 4 and 
612 2 have been deposited in ZMJU. 

Other material examined: Berlese Acaroteca (EIAZ), 19 (slide 6/40), 49 2 (6/41), 
288,222 (7/37), 584,299, 2 deutonymphs (7/35). All this material was collected in 
Vallombrosa, but the habitat was not specified. 

Diagnosis. The species is recognisable by very thick seta pv! on tibia IV in both 
sexes (Fig. 2 A,B). 

Description. Male. Idiosoma brown, well sclerotized. Dimensions of idiosoma: 
555-585 x 365-380 um; L/W (length/width) factor 1.46-1.56, N=10. Podonotal region 
with 20 pairs of setae, j1 = 38-40 um, other 19-22 um; opisthonotal region with 30 
pairs of shorter setae, 10-13 um. 

Ventral side (Fig. 2 C). Genital lamina large, located in a well sclerotized 
concavity, its lateral sides forming triangular sharp projections, its anterior edge with 
median pleated membrane and two lobes; postero-lateral edges of genital lamina with 
two well sclerotized protrusions. 

Male genital orifice, flanked by triangular presternal shields and provided with 
large rectangular microsclerite bearing tritosternum. Reticulation of sternogenital 
region regular; between anterior margin and close to sternal setae 1 (stl), sternal shield 
more heavily sclerotized and its reticulation slightly convex; length of sternal setae 
about 42 um. Opisthogastric region with 8 pairs of setae (26-39 um). 

Gnathosoma. Tectum trispinate, with long, broad central prong (Fig. 3 I). Corni- 
culi with paraxial margin forming rounded lamellar protrusion (Fig. 2 D,E). Hypo- 
gnathal groove with 11 complete rows of denticles; palpcoxal setae pilose, hypo- 
stomatic setae simple (Fig. 2 D). Chelicera (Fig. 2 F,G). Both digits short and robust. 
Fixed digit toothless, with pilus dentilis flanked by two convex laminae. Movable digit 
with two teeth. Arthrodial membrane at the base of movable digit with well developed 


NEW SPECIES OF HOLOPARASITUS 9 


Fic. 2 


Holoparasitus crassisetosus sp. n. Male: A-tibia IV; C-sternogenital region and genital lamina; 
D-gnathosoma, palptrochanter and palpfemur; ventral; E-corniculus; F-chelicera, antiaxial: 
G-idem, paraxial; H-leg II, femur, genu and tibia, antiaxial; I-tibia, ventral; J-basis tarsus and 
cuticular edge of tibia; Female: B-tibia IV. 


10 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


brush-like process on paraxial side and a lesser developed one on antiaxial side, 
synarthrodial membrane short and wedge-shaped. 

Pedipalps. Trochanter with proximal seta (v1) simple and distal seta (v2) pilose. 
Femur with tubercle located on ventral side close to anterolateral seta. Anterolateral 
seta of femur spatulate and pectinate on inner side (Fig. 2 D); anterolateral setae of 
genu spatulate. 

Legs. Coxae II with ridge of 5 denticles (Fig. 3 J). Leg II armed as shown in 
figure 2 H-I. Femoral apophysis finger-shaped and straight, axillary process curved 
towards the femur. Spur on genu small, round, located ventrally in the middle. 
Apophysis on tibia low and long, with slightly convex margin, attached ventrally and 
reaching with its rounded distal end the margin of segment; the cuticle of anterodistal 
edge of tibia with several fine furrows. Trochanter IV with flattened protuberance 
situated medioposteriorly (Fig. 2 K). On tibia IV seta pv! conspicuous, very thick with 
a dentate end (Fig. 2 A). Measurements: tarsus I = 139-144 um; tarsus IV = 146um. 

Female. Idiosoma brown, well sclerotized. Dimensions of idiosoma: 630-665 x 
445-475um (L/W factor 1.36-1.45, N=10). Podonotal setae: j1 = 39 um, other setae 19- 
26um; opisthonotal setae 12-18 um. 

Ventral side (Fig. 3 A). Fused presternal shields forming smooth ribbon, 
partially connected to lateral platelets. Anterior margin of sternal plate sometimes with 
incisions of a soft cuticle, laterally to setae stl. Reticulation of sternal shield with two 
prominent lines delimiting a slightly more sclerotized anterior region of sternum: one 
line V-shaped extending medially and second line running close to second pair of pores 
(pst2). Margin of exopodal plate facing coxa I sclerotized. 

Genital region. Epigynial plate heptagonal, anterior margin with a middle 
triangular apex and two lateral spines curved antiaxially; subapical structure small and 
oval, with lateral rounded protrusions extending beyond epigynial margin (Fig. 3 C). 
Paragynia rectangular with elliptical thickenings in front of coxae IV and not extending 
beyond paragynium edge, paragynial posterior protrusions elongated (Fig. 3 B). 
Endogynium oval, 22 x 29 to 24 x 31 um. Distance between endogynial margin and 
coxa IV nearly two times larger than endogynium length (56-59 um). Shape of 
endogynium cup-like, with anterior margin protruding into dorsally directed appendage, 
which looks like hollow tube (Fig. 3 D). Gland pores behind coxa IV large. Opistho- 
gastric region with 8 pairs of setae, their length 36-39 um. 

Gnathosoma. Tectum trispinate, central prong long and narrow, lateral prongs 
small; minute additional spines discernible between central and lateral prongs (Fig. 
3 E). Corniculi conical. Hypognathal groove with 6-8 rows of denticles ending at 
palpcoxal setae level; some lateral lines present, starting from hypognathal groove; 
cuticular ridges between palpcoxal setae serrated (Fig. 3 F). Palpcoxal setae slightly 
pilose, hypostomatic setae simple. 

Chelicera. Fixed digit with two teeth in front of pilus dentilis, a third, smaller 
tooth frequently present between them; two teeth and a lamella with slightly concave 
edge located behind pilus dentilis. Movable digit with three teeth (Fig. 3 G). 

Pedipalps. Proximal seta of trochanter (v1) simple, distal seta (v2) pilose (Fig. 3 
F), anterolateral seta of femur spatulate and pectinate on one side, and anterolateral 
setae of genu spatulate. Femur with poorly visible tubercle located anterolaterally. 


NEW SPECIES OF HOLOPARASITUS 11 


FIG. 3 


Holoparasitus crassisetosus sp. n. Female: A-presternal and sternal shields; B-paragynium; C- 
epigynium; D-endogynium; E-tectum; F-gnathosoma and palptrochanter, ventral; G-chelicera, 
paraxial; H-trochanter IV, ventral. Male: I-tectum; J-coxa II, denticulated ridge. 


12 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


Legs. Coxa II with a ridge of about 5 denticles on anterior face. Tibia IV with 
very thick pv1 seta, its end dentate (Fig. 2 B). Trochanter IV with rounded protuberance 
located on proximal third of posterior face (Fig. 3 H). 

Measurements: tarsus I = 144-154 um; tarsus IV = 156-163 um. 

Discussion. Males and females of H. crassisetosus are very similar to those of 
H. digitiformis. From other Holoparasitus species known to date these species differ in 
the following features. In males, excipulum is absent, tectum trispinate, movable and 
fixed digits of chelicera with 2 and 0 teeth, respectively. Corniculi posses a lamellar 
dilatation or indentation on the paraxial margin. In females, presternal plates and lateral 
platelets accreted, the endogynium soup-spoon-shaped. The most easily recognized 
feature which distinguishes H. crassisetosus and H. digitiformis is the presence of the 
transformed pvl seta of tibia IV in both sexes of the former species. A detailed 
discussion of the most important differences between the two species is given at the end 
of the description of H. digitiformis. 


Holoparasitus digitiformis sp. n. Figs 4, 5 


Type material: 15 holotype, 526 8, 392 © paratypes, from litter of oak forest (Quercus 
cerris, Q. pubescens) and 68 6d and 729 2 paratypes, from litter of pine forest (Pinus nigra, 
Erica arborea, Juniperus communis). The two forests are situated at Selvapiano (Commune di 
Rufina, Toscana), 25 km away from Florence at an altitude of 200 m. The two habitats are about 
300 m apart and are situated on the same side of the mountain, facing south-west. The material 
was collected from February 1971 to April 1974 by Dr. R. Nannelli and Dr. F. Pegazzano 
(EIAZ). 

Other material examined: 193 4, 159 ?, Catena del Marghine (Marghine Range), 
Sardinia, from moss and lichens in yew (Taxus baccata) and oak forests along the road to 
Bolotana, alt. ca 1000 m, 30.03.1977, collected by the staff of the DEBS and deposited in ZMJU. 

12 (slide 6/42), 18 (6/43), 12 (6/44), Monte Senario, Toscana, Berlese Acaroteca 
(EIAZ). The material has been deposited as follows: 1d holotype, 326 d, 199 © paratypes from 
oak forest and 48d d, 529 © paratypes from pine forest — in MHNG; 203 d, 209 © paratypes 
from both habitats - in ZMJU; 203 4,209 © paratypes from both habitats - in EIAZ. 

Diagnosis. In both sexes all setae on tibia IV of normal appearance. Male: 
excipulum absent; movable digit of chelicera with two denticles, fixed digit edentate, 
slender, finger-like, with slightly dilated apex. Female: epigynium with subapical small 
ovoid microsclerite and hyaline wing-like protrusions extending anterolaterally (Fig. 5 
D). Endogynium small, oval, inverted cup-like, with anterior margin protruding into a 
cuticular solid appendage directed more or less dorsally (Fig. 5 B,C,E). 

Description. Male. Idiosoma well sclerotized. Dimensions of idiosoma: speci- 
mens from Selvapiano, Toscana, oak forest: length 576-617 um; idem, specimens from 
pine wood, length 634-641 um. Specimens from Sardinia, idiosoma size 585-635 x 
405-430 um (L/W factor 1.45-1.57, N=10). Podonotal region with 21 pairs of seta, 
j1=39 um, the others 26-28 um opisthonotal region with about 30 pairs, 19-24 um. 

Ventral side. Genital lamina large, situated in well defined concavity of heavily 
sclerotized anterior margin of sternal shield; anterior margin of the lamina with two fine 
lobes separated by a concavity and two lateral triangular prongs. Between lateral prongs 
and anterior lobes, on their inner face, two well sclerotized prominences. Behind genital 
lamina a rectangular microsclerite with a rounded lobe ventrally supporting the base of 


NEW SPECIES OF HOLOPARASITUS 


Fic. 4 


Holoparasitus digitiformis sp. n. Male: A-genital lamina; B-sternogenital shield; C-tectum; 
gnathosoma and palptrochanter, palpfemur, ventral; E-chelicera; F-idem, paraxial; G-leg 
femur, genu and tibia, paraxial; H-femoral apophysis, antiaxial; I-trochanter IV, ventral. 


ID: 
II, 


14 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


tritosternum (Fig. 4 A). Sternogenital shield with scale-like reticulation and with 
marked line (inverted V) behind second pair of sternal setae; length of sternal setae 39- 
46 um (Fig. 4 B). Large gland pore behind coxae IV. On opisthogaster 8 pairs of setae, 
24-36 um. 

Gnathosoma. Tectum trispinate, lateral prongs small, the central one large, broad 
at the base and pointed (Fig. 4 C). Corniculi with small prominence situated paraxially; 
hypognathal groove with 10-11 rows of denticles; palpcoxal setae finely pilose, hypo- 
stomatic setae simple (Fig. 4 D). 

Chelicera (Fig. 4 E,F). Fixed digit edentate, slender, finger-like; its apex slightly 
dilated and sometimes curved. Movable digit with two denticles, the distal one 
sometimes appearing much more paraxially, synarthrodial membrane rounded. Sperma- 
todactyl large, with a median concavity; arthrodial membrane with short brushy process 
paraxially and setiform margin antiaxially. 

Pedipalps. Palptrochanter with seta vl thin and simple, whereas v2 thicker and 
pilose. Palpfemur with anterolateral seta pectinate on one edge; palpgenual anterolateral 
setae simply spatulate. Palpfemoral segment swollen distally, with tubercle located on 
ventral side, close to al seta (Fig. 4 D). 

Legs. Coxa I with a ridge bearing 6-7 denticles, situated paraxially close to 
distal margin. Coxa II with ridge of 8-10 denticles (Fig. 5 J). Armature of leg II illus- 
trated in figure 4 G. 

Femoral apophysis relatively long, finger-shaped, with curved apex and a small 
tubercle on its base (Fig. 4 H); axillary process curved. Genu with small rounded spur 
situated medioventrally. Tibia with two spurs: one low and elongated, reaching distal 
margin, another smaller, located anterolaterally and extending beyond segment margin. 
Posterior face of trochanter IV with a small tubercle under seta pv (Fig. 4 I). 

Measurements: specimens from Toscana: tarsus I = 156-158 um; tarsus IV = 
168-173 um. 

Female. Idiosoma well sclerotized, brown coloured. Dimensions of idiosoma: 
660-710 x 490-530 um (L/W factor 1.28-1.42, N=10). Podonotal region with 21 pairs 
of setae, }1=36um, the others 16-20 um; opisthonotal region with 33 pairs of short setae 
(13-16 um). 

Ventral side. Accreted presternal shields forming a smooth ribbon-shaped plate, 
more narrow medially; lateral presternal platelets usually free. Anterior margin of 
sternal plate with two less sclerotized concavities, frequently flanking bases of setae 
stl; reticulation of sternum easily discernible with prominent arched transverse line 
passing close to pores pst2 (Fig. 5 A); length of sternal setae from 46 um (stl) to 65 um 
(st3). Anterior edge of exopodal shield facing leg I thickened. 

Genital region. Paragynial shield with sclerotized elliptical thickening in front of 
coxa IV (Fig. 5 B.C). Postero-lateral protrusions long and narrow, ending with rounded 
or sometimes pointed edge. Metagynial sclerites with paraxial margins convex. 
Epigynial shield with anterior margin formed by a triangular central apex and two large 
lateral prongs curved antiaxially; subapical epigynial structure with small ovoid 
microsclerite and hyaline wing-like protrusions extending anterolaterally (Fig. 5 D). 
Endogynium (Fig. 5 B,C,E) small, oval (20 x 27 um to 27 x 33 um), inverted cup-like, 


NEW SPECIES OF HOLOPARASITUS 15 


li 0,1mm. A,B,C,D,J 
SU Ma 
me “i 005mm. EI 
Fic. 5 


Holoparasitus digitiformis sp. n. Female.: A-presternal and sternal shields; B-paragynia and 
endogynium, dorsal; C-idem, ventral; D-epigynium; E-endogynium; F-tectum; G-hypognathal 
groove and corniculus; H-chelicera, antiaxial; I-coxa II, denticulated ridge. Male: J-coxa II, 
paraxial. 


16 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


with anterior margin protruding into a cuticular solid appendage directed more or less 
dorsally; poorly visible scale-like fine hyaline flap covering endogynium on ventral 
side. Gland pores behind coxa IV large. Opisthogastric region with 8 pairs of setae, 
their length about 39 um. 

Gnathosoma. Tectum usually trispinate, with central prong sharply pointed, 
sometimes an extra spine present on one side (Fig. 5 F). Corniculi conical. Hypognathal 
groove with 8-11 rows of denticles, those located posterior to the level of palpcoxal 
setae frequently reduced or absent, some lateral lines present between hypostomatics 
and palpcoxal setae (Fig. 5 G ). Palpcoxal setae pilose, hypostomatics simple. 

Chelicera. Fixed digit with two teeth in front of pilus dentilis, two more teeth 
and a thin, concave cuticular edge behind pilus dentilis. Movable digit with three teeth 
(Fig. 5 H). 

Pedipalps. Trochanter ventral setae finely pilose; anterolateral seta of femur 
spatulate and pectinate on one edge; anterolateral setae of genu simply spatulate. Palp- 
femur swollen distally, with tubercle located on ventral side close to anterolateral seta. 

Legs. Leg structure and setation unremarkable. Coxa I with row of about 10 
little denticles on distal paraxial margin. Coxa II with ridge of 8 to 10 fine denticles 
situated on anterior face (Fig. 5 I). Measurements: tarsus I = 154-163 um; tarsus IV = 
168-175 um. 

Discussion. H. digitiformis is similar to H. crassisetosus in respect to the general 
structure of the genital region in the female and to the shape of the armature of leg II in 
the male. 

The endogynium in both species is small and inverted cup-shaped, with the 
anterior margin forming a cuticular protrusion directed dorsally, thus similar in 
appearance to a soup-spoon. In H. crassisetosus, this protrusion is relatively shorter (as 
compared with the size of endogynium) than in H. digitiformis and appears as a hollow 
tube. The metagynial sclerites are oriented roughly parallel in both species, but in A. 
digitiformis their lateral margins are convex, whereas in H. crassisetosus they are rather 
concave. The subapical epigynial structure has prominent wing-like protrusions in H. 
digitiformis but only small round protrusions in H. crassisetosus . 

Males in both species have a very similar armature of leg II, especially on the 
femur and genu: on the tibia, however, in A. digitiformis the additional prominent 
anterolateral tubercle is present, whereas in H. crassisetosus the cuticle of corres- 
ponding place shows fine furrows only. A very pronounced difference can be seen in 
the shape of the male corniculi: H. digitiformis has a small protuberance on the paraxial 
margin, whereas in H. crassisetosus this margin extends prominently forming a thin 
rounded lamella. 

In both sexes, the most important character for differentiation of these two 
species remains the form of seta pv! of tibia IV: it is narrow and simple in H. digi- 
tiformis but thick and notched at the end in A. crassisetosus. H. digitiformis was found 
in the surroundings of Florence (Toscana) in coniferous forest as well as in Sardinia 
where it occurs in soil of yew and oak forests. H. crassisetosus was collected, till now, 
only from Vallombrosa (Toscana), where it lives in coniferous forests. 


NEW SPECIES OF HOLOPARASITUS 17 


TAR 


0,05mm. 


Fic. 6 


Holoparasitus cornutus sp.n. Male: A-scutum sternogenital and genital lamina; B-tectum: 
C-tectum (Bevagna-Umbria, Berlese Acaroteca); D-gnathosoma and palptrochanter, ventral; 
E-chelicera, antiaxial; F-leg II, femur, genu and tibia; G-tibia II, ventral (Bevagna-Umbria, 
Berlese Acaroteca); H-coxa II, denticulate ridge; I-trochanter IV, ventral; J-idem, dorsal. 


18 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


Holoparasitus cornutus sp. n. Figs. 6, 7 


H. apenninorum sensu Juvara-Bals 1975: 400: nec Berlese, 1906. 

Type material: 13 holotype, 26 4, 42 2 paratypes, Vallombrosa (Toscana, Italy), litter 
in beach forest, 19.09.1971: 148 &, 119 2 paratypes, Vallombrosa, moss in coniferous forest 
(Abies alba), alt. 960 m, 6.11.1982. All paratypes collected by the staff of DEBS. The holotype 
and 14,29 2, 2 deutonymphs have been deposited in MHNG, whereas 166 6, 132 © in ZMJU. 

Other material examined. Italy: 18, 12, Monte Amiata near Siena, Toscana, litter in 
beach forest, alt. ca. 900 m; 49 9, Monte Amiata, litter in chestnut forest, alt. ca. 800 m, 
17.09.1990, leg. W. Witalinski, deposited in ZMJU. 16, 32 © (slide 7/40), Monte alle Forche, 
Toscana, and 15,1% (slide 7/39), Bevagna, Umbria, Berlese Acaroteca (EIAZ). 

Romania: 16, 19, Cerbului Valley, Busteni, Bucegi Mountain, Meridional Carpathians, 
sawdust and litter, beech and spruce forest, 5.6.1967, leg. I. Juvara-Bals, deposited in MHNG. 

Diagnosis. The species can be recognized by a lamellar process ending with one 
or two spurs, situated more or less dorsally at the distal margin of trochanter IV in both 
sexes. Male: excipulum absent; two incisions in sclerotized cuticle behind each group 
of hypostomatic setae; folded cuticle at the level of legs IV. Female: endogynial sack 
rounded, with two large, connected teeth on each side . 

Description. Male. Idiosoma well sclerotized, its dimensions (Italy) 672-715 x 
452-510 um (L/W factor 1.42-1.48, N=5). Podonotal setae: j1=38 um, other setae 20- 
26 um; opisthonotal setae shorter, 13-19 um. 

Ventral side (Fig. 6 A). Genital lamina with serrated anterior margin. Sternal 
shield reticulated with a slight median prominence distally; two ridges run from this 
prominence posteriorly to sternal setae st2 towards the margin of the shield. At the level 
of coxa IV, cuticle folded in a strongly sclerotized line. Length of sternal setae from 48 
um (stl) to 40 um (st3). On opisthogastric region 8 pairs of setae, their length from 42 
um (Jv1) to 26 um (Jv5). 

Gnathosoma. Tectum trispinate with central prong triangular, long, and in the 
specimen from Bevagna, with a little denticle on its tip; lateral prongs tiny (Fig. 6 B,C). 
Hypognathal groove provided with 11 well dentated rows; gnathosomal setae pilose, 
except simple hypostomatic seta 3. Behind hypostomatic setae there are incisions in 
sclerotized cuticle (Fig. 6 D). Corniculi conical. Chelicera (Fig. 6 E). Fixed digit eden- 
tate and with truncate apex; movable digit with 2-4 small subapical teeth. Arthrodial 
membrane formed by a short brush-like processes paraxially and a setiform fringe 
antiaxially. 

Pedipalps. Palptrochanter with seta vl simple and v2 pilose; palpfemur with 
distal protuberance on its ventral face. 

Legs. Coxa II with ridge bearing 9 denticles located on anterior face (Fig. 6 H). 
Armature of leg II shown in figure 6 F. Main femoral apophysis long, finger-like, its 
axillary spur rounded. Spur on genu rounded. Tibia with median saddle-like spur and 
little distal protuberance situated anterolaterally (Fig. 6 F,G). Dorsal or posterodorsal 
face of trochanter IV with distal lamellar process ending with one or two spurs; 
proximal posterolateral face with a protuberance (Fig. 6 I,J). Gland pores behind coxae 
IV located in a normal cuticle. Measurements: tarsus I = 143 um (specimen from 
Romania) and 173-177 um (specimens from Italy); tarsus IV = 173 um (Romania) and 
184-193 um (Italy). 


NEW SPECIES OF HOLOPARASITUS 19 


0,05mm. 


le] 


FIG.7 


Holoparasitus cornutus sp. n. Female: A-presternal and sternal shields; B-paragynium; C- 
epigynium; D-endogynium (2 from Rumania); E-idem, (9 from Italy); F-tectum; G-hypognathal 
groove and corniculi; H-palptrochanter and palpfemur, antiaxial; I-chelicera, paraxial; J-tro- 
chanter IV, ventral; K-coxa II, denticulate ridge; L-gland gv2. 


20 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


Female. Dimensions of idiosoma: 580 x 430 um (L/W factor 1.35), specimen 
from Romania; 742-806 x 550-580 um (L/W factor 1.34-1.41, N=5), specimens from 
Italy. Podonotal setae from 13 um to 39 um, j1 = 36-38 um; opisthonotal setae shorter, 
about 13 um. ; 

Ventral side. Presternal shield smooth, ribbon-like, with two lateral excres- 
cences, sternal shield with transverse ridge at level of pst2, some specimens with two 
lightly sclerotized indentations in front of setae stl (Fig. 7 A). Sternal setae slender, 
55-60 um. Paragynial shield provided on its dorsal face with an elliptical thickening in 
front of coxa IV and on its inner ventral margin with one or two denticles; metagynial 
sclerite elongated, paragynial posterior protrusion elongated and rounded, paragynial 
lobe trapezoidal (Fig. 7 B). Epigynium heptagonal; its apex transparent, its subapical 
structure formed by a fine rounded margin and a sclerotized three-lobed thickening 
(Fig. 7 C). Endogynium: a rounded sack with two large, connected teeth on each side, 
covered by a scale flap ventrally, sometimes provided with denticles on inner walls; 
dorsally, its cuticle with many pores arranged in ca. 8 rows of muscle-prints (Fig. 7 
D,E.). Gland pores behind each coxa IV located in a normal cuticle (Fig. 7 L). On 
opisthogaster 8 pairs of setae, their length from 40 um (Jv1) to 24 um (Jv5). 

Gnathosoma. Tectum trispinate, lateral prongs tiny (Fig. 7 F). Hypognathal 
groove with 9-10 lines, anterior dentate, posterior bare; external posterior hypostomatic 
seta simple, other gnathosomal setae slightly pilose. Corniculi conical (Fig. 7 G). 

Chelicera. Fixed digit with 3 minute denticles in front of pilus dentilis and 
behind it two teeth and a cuticular edge (Fig. 7 I). 

Pedipalps. Trochanter with seta vl simple and seta v2 slightly pilose. Femur 
with small protuberance near al seta (Fig. 7 H). 

Legs. Coxa IT with ridge bearing about 6 denticles (Fig. 7 K). Trochanter IV 
provided proximally with rounded protuberance on posterolateral face and with lamel- 
lar process, pointed in one or two spurs, dorsally or posterodorsally on distal margin of 
segment (Fig. 7 J). Measurements: tarsus I = 145 um (specimen from Romania) and 
164-173 um (specimens from Italy); tarsus IV = 166 um (Romania) and 189-196 um 
(Italy). 

Discussion. Juvara-Bals (1975) identified and described under the name of 
H. apenninorum (Berlese) specimens of Holoparasitus collected in the surroundings of 
Siena (Italy) and in Bucegi Mountain, Meridional Carpathians (Romania). She thought 
that the material belonged to H. apenninorum according to characteristics of male 
chelicera. In fact, Berlese (1906) had drawn two types of chelicera for O. pollicipatus 
var. apenninorum, as already discussed in the comments to the redescription of that 
species. Figure 16, tab. XIII by Berlese (1906) was a drawing of a specimen from 
Monte Senario which is the same as H. cornutus, the new species described above. The 
specimens of A. cornutus from Romania and Italy do not exhibit significant differences 
(Juvara-Bals, 1975). The male genital lamina is not serrated in specimens from Roma- 
nia but instead it has only two central denticles. The number of teeth on the movable 
digit is 2 in Romanian versus 3 in Italian specimens and the shape of the tectum is 
slightly variable. H. cornutus is characterised especially by the lamellar process situated 
more or less dorsally on margin of trochanter IV in both sexes. Other characteristics 


NEW SPECIES OF HOLOPARASITUS 2] 


FIG. 8 


Holoparasitus gibber sp. n. Male: A-chelicera, paraxial; B-idem, antiaxial; C-base of movable 
digit; D-tectum; E-gnathosoma and palptrochanter, ventral; F-corniculus; G-palptrochanter and 
palpfemur; H-genital lamina; I-coxa I, paraxial; J-coxa II, denticulate ridge; K-leg II, femur, genu 
and tibia. 


22 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


such as the shape of the chelicera and armature of leg H in the male, as well as the 
particularities of the endo- and epigynium of the female distinguish this species from 
the other species belonging to the complex O. pollicipatus var. apenninorum sensu 
Berlese 1906. We also have to mention the species H. hemisphaericus (Vitzthum, 1923) 
described on the basis of a single female found in Austria. This species was considered 
by Juvara-Bals (1975) to be synonymous with H. apenninorum sensu Juvara-Bals, 
1975. H. cornutus was compared with the type of H. hemisphaericus kindly sent to the 
senior author by Dr. L. Tiefenbacher (Munich, Germany). H. hemisphaericus is a valid 
species which differs from H. cornutus by the shape and size of the dorsal protuberance 
located distally on trochanter IV, by the absence of the proximal protuberance on 
trochanter IV, as well as by the paragynia, characterised by thickenings at paraxial 
margins but devoid of any thickening in front of coxa IV. Unfortunately, this species is 
known only on the basis of one female and on the single specimen available the 
characteristics of the epigynium and those of the endogynium cannot be properly 
observed. 


Holoparasitus gibber sp. n. Figs 8, 9 

The species is described from permanent slides of the Athias-Henriot collection. The 
length of the idiosoma cannot be measured because the specimens are crushed. 

Type material: 13 holotype, 26 4, 32 2, 1 deutonymph, paratypes, Los Palacios (near 
Seville, Spain), sifting of flood litter, 22.02.1951, leg. H. Franz, and 48 8,4? 2, 1 deutonymph 
paratypes, surroundings of Carmona (near Seville, Spain), in the vicinity of Finca Alamaja, black 
soil from decayed plants, 22.02.1951, leg. H. Franz. All types in collection of Athias-Henriot 
deposited in MHNG. 

Diagnosis. Male: excipulum absent; mobile digit of chelicera with | tooth, fixed 
digit with subapically located 6 denticles plus one (Fig. 8 A,B); hypostome extended 
between corniculi. Female: presternal plate with anterior margin serrated; endogynium 
circular in outline with two prolongations of posterior margin reaching anterior edge of 
endogynium, lateral sides of endogynial sack with 1-2 denticles (Fig. 9 D). 

Description. Male. Idiosoma. Cuticle yellow-brownish, moderately sclerotized. 
Setae on holodorsal shield from 36 um (j1) to 13-15 um in opisthonotal region. 

Ventral side. Sternal shield reticulated, without a particular pattern; length of 
sternal setae 24 um (st2) to about 36 um (stl, st4). Genital lamina with large central 
rounded process (Fig. 8 H). Opisthogaster with 8-9 pairs of setae, their lengths about 
20 um. Simple gland pore behind coxae IV. 

Gnathosoma. Tectum with three prongs, the median much larger and broader 
than lateral ones (Fig. 8 D). Hypostome extended between corniculi; hypognathal 
groove provided with 9 rows of very fine denticles: hypostomatic and palpcoxal setae 
simple. Sclerotized cuticle with incisions behind hypostomatics; internal malae covered 
by a trapezoidal extension (Fig. $ E). Corniculi slender and conical (Fig. 8 F). 

Chelicera (Fig. 8 A-C). Fixed digit straight and narrow, its apex slightly curved; 
along its upper third, internal paraxial edge provided with 6 small denticles and 
antiaxial edge only with one big tooth; pilus dentilis located between these two edges. 
Movable digit with big tooth medially on internal margin and a characteristic rounded 


NEW SPECIES OF HOLOPARASITUS 


(NO) 
Ww 


hump on external (ventral) margin. Spermatotreme slender, ending distally at the level 
of tooth: the basal part of the digit with well sclerotized enlargement and a little 
denticle. Arthrodial membrane developed into brush-like process. 

Pedipalps. Trochanter with protuberance located between slightly pilose setae v1 
and thick and pilose v2; femur provided with small rounded protuberance, seta al 
slightly spatulate (Fig. 8 G). 

Legs. Distal margin of coxa I dentate, coxa II with ridge bearing 7-8 denticles 
and an extra basal denticle (Fig. 8 I,J). Spurs on leg II as in figure 8 K: femoral 
apophysis short and thumb-like and axillary process trapezoidal; triangular spur located 
distally on genual margin; tibial apophysis elongated, situated medially on anterolateral 
face. Measurements: tarsus I = 159-169 um; tarsus IV = 164-169 um. 

Female. Idiosoma. Colour yellow-brownish; setae on podonotal region from 46 
um (j1) to 26 um; setae on opisthonotal region shorter, ca. 13 um. 

Ventral side. Presternal plate entire, ribbon-like, with the anterior margin 
serrated; lateral presternal platelets free. Sternal shield reticulated with a longitudinal 
granular strip medially (Fig. 9 A), length of sternal setae about 52 um. Paragynial 
shields reticulated, metagynial sclerites oval (Fig. 9 B). Epigynium heptagonal, its 
anterior margin with triangular median apex and two lateral prominent spines: 
subapical epigynial structure formed by a weakly sclerotized rectangle and a hyaline 
structure stretching beyond epigynial margin in the form of two little wings (Fig. 9 C). 
Endogynium cup-shaped and circular in outline. Its posterior margin protrudes forward 
to form two long, sinuous prolongations with rounded tips reaching anterior edge of 
endogynial sack; lateral sides with 1-2 denticles (Fig. 9 D). Opisthogastric shield with 9 
pairs of setae, their length from 42 um (Zv1) to 24 um (Jv5). 

Gnathosoma. Tectum trispinate with long median prong and two tiny lateral 
prongs; dorsal face slightly granular (Fig. 9 E). Hypognathal groove with 9-10 rows of 
denticles, last four oligodentate; palpcoxal seta pilose, anterior hypostomatic seta 
slightly barbed, posterior setae simple (Fig. 9 F). 

Pedipalps (Fig. 9 G). Border of trochanter thickened between pilose setae vl and 
v2; distal margin of femur with small tubercle. Chelicera (Fig. 9 H) similar to that of H. 
ellipticus: fixed digit with five teeth and a long, thin cuticular ridge, movable digit with 
three teeth. 

Legs. Structure and setation unremarkable. Denticulate ridge on coxa II as in the 
male (Fig. 9 I). Measurements: tarsus I = 156-163 um; tarsus IV = 173-185 um. 

Discussion. Adults of H. gibber are similar to those of H. mallorcae and of 
H. ellipticus. All these species have in common: denticulation of the anterior margin of 
the female’s presternal shield, the shape of the endogynium and the pattern of male’s 
second leg armature; but they differ mainly in the shape of the male’s chelicera and 
tectum as well as in the ferm of the female’s epigynium. See comments following the 
description of H. ellipticus. 


Holoparasitus ellipticus sp. n. Fos OM 


Type material: 15 holotype, 116 8,9% © paratypes, Milazzo, Sicily, Italy from litter of 
olive trees, 24.03.1972; 18, 29 2 paratypes, Monti Peloritani, Sicily, from humus and litter in 


24 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


Fic. 9 


Holoparasitus gibber sp. n. Female: A-presternal and sternal shields; B-paragynium; C-epigy- 
nium; D-endogynium; E-tectum; F-hypognathal groove and corniculus; G-palptrochanter and 
palpfemur: H-chelicera, paraxial; I-coxa II, denticulate ridge. 


oak forest, 26.03.1972. The material was collected by the staff of the DEBS. The holotype and 
36 d, 69 9 have been deposited in MHNG, whereas 98 8,52 2 in ZMJU. 


Diagnosis. Male: excipulum absent; mobile and fixed digits of chelicera with 
many denticles (Fig. 10 D,E); hypostome evidently extended between corniculi. 


NEW SPECIES OF HOLOPARASITUS DS 


Fic. 10 


Holoparasitus ellipticus sp. n. Male: A-genital lamina and sternogenital shields; B-tectum; C- 
gnathosoma and corniculi; D-chelicera, paraxial; E-idem, antiaxial; F-palptrochanter and 
palpfemur; G-G’-leg II, femur, genu and tibia antiaxial and ventral; H-coxa II, paraxial. 


26 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


Female: presternal shield with anterior margin serrated; endogynium circular in outline; 
its posterior margin protrudes forward to form two prolongations ending beyond the 
anterior margin of endogynial sack; endogynial margin and walls with 3-7 denticles 
(Fig. 11 D). 

Description. Male. Idiosoma markedly elongated, moderately sclerotized. 
Dimensions of idiosoma: 615-655 x 390-410 um (L/W factor 1.57-1.64, N=10). Length 
of setae: podonotal region 15-20 um, jl=39 um and opisthonotal region 10-13 um. 

Ventral side. Genital opening located in a shallow concavity of sternal shield 
margin. Genital lamina divided into three lobes: central one bilobate, protruding for- 
ward more than lateral ones (Fig. 10 A). Sternogenital region with regular reticulation; 
length of sternal setae 39 um. Cuticle around gland pores behind coxae IV not 
modified. 

Gnathosoma. Tectum trispinate, with all lateral prongs terminated with minute 
spines; central prong rounded (Fig. 10 B). Hypostome with conspicuous broad exten- 
sion located between slender corniculi. Hypognathal groove with 10 rows of denticles. 
Palpcoxal setae pilose, hypostomatic setae simple. Incisions in sclerotized cuticle 
behind hypostomatic setae (Fig.10 C). 

Chelicera (Fig. 10 D,E). Fixed digit slender; its proximal 3/5 toothless, with 
parallel edges; distal part equipped with 4-5 small denticles paraxially, and pilus 
dentilis inserted in the vicinity of two more or less pronounced tubercles. Movable digit 
curved, bearing 5-7 denticles distally and large tooth medially; dorsal face of digit 
weakly sclerotized proximally. An elongated protuberance located paraxially at the 
base of spermatotreme. Arthrodial membrane with brush-like process developed 
paraxially, synarthrodial membrane rounded. 

Pedipalps (Fig. 10 F). Paraxial border of trochanter slightly thickened, visible as 
a ridge, ending near seta vl; seta vl thin and barbed along one edge, whereas seta v2 
thick and richly pilose. Ventral surface of femur convex. 

Legs. Legs I, IN and IV unremarkable. Leg II spurred as follows (Fig. 10 G,G’): 
main spur on femur relatively short and rounded, axillary spur rounded and located 
close to main spur, both spurs end at the same level. Genu II with conical spur located 
distally and extending slightly beyond distal margin. Spur on tibia longer than that on 
genu, situated more paraxially and attached to tibia surface over a long distance. Ridge 
of 8 denticles and one solitary denticle on coxa II (Fig. 10 H). Measurements: tarsus I = 
160-168 um; tarsus IV = 173-184 um. 

Female. Idiosoma elongated but highly convex, cuticle moderately sclerotized, 
yellow to yellowish-brown. Dimensions of idiosoma: 690-770 x 465-540 um (L/W 
factor 1.43-1.52, N=10). Podonotal setae: j1 = 45-46 um, the others 20-39 um; opistho- 
notal setae shorter, 10-14 um. 

Ventral side. Presternal plates entire, ribbon-like, with denticulate anterior 
margin; lateral presternal platelets free. Exopodal shields facing legs I with thickened 
anterior edge and connected to sternum via thin cuticle. Reticulation of sternum weakly 
pronounced, more visible in anterior region; posterior region with an axially running 
granulate strip (Fig. 11 A). Length of sternal setae from 52 um to 59 um. 


NEW SPECIES OF HOLOPARASITUS 27 


Fıc. 11 


Holoparasitus ellipticus sp. n. Female: A-presternal and sternal shields; B-paragynium: C-epi- 
gynium; D-endogynium; E-tectum; F-hypognathal groove and corniculus; G-palptrochanter: 
H- chelicera, paraxial; I-gland gv2. 


Epigynium characterised by sharp lateral spines separated from central apex by 
deep concavities; lateral protrusions of subapical epigynial structure extending slightly 
beyond epigynium margin (Fig. 11 C). Paragynial shields with oval and slightly 
concave metagynial sclerites; thickenings in front of coxae IV absent. Paragynial 


28 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


protrusions “locking” epigynial shield variable, ranging from ones with border 
paraxially angular to circular ones (Fig. 11 B). Cup-shaped endogynium circular; its 
posterior margin protrudes forward to form two slightly sinuous, tongue-like ribbons 
ending beyond anterior margin of endogynium; endogynial margin and walls armed 
with several (3-7) denticles, distributed more or less regularly. Endogynial opening 
covered with delicate lamina (Fig. 11 D). Opisthogaster with 8 pairs of setae, their 
length from 36 um to 48 um. Large, singular gland pores behind coxa IV surrounded by 
unmodified cuticle (Fig. 11 I). 

Gnathosoma. Tectum trispinate, with long, attenuate median tine (Fig. 11 E). 
Hypognathal groove with 10 rows of denticles. Palpcoxal setae pilose, hypostomatic 
setae simple, corniculi conical (Fig. 11 F). 

Chelicera (Fig. 11 H). Fixed digit with three teeth in front of pilus dentilis; 
sometimes central one smaller and located more paraxially. Pilus dentilis protected 
partially by truncated cuticular elevation; two teeth and a long, thin cuticular ridge 
located behind pilus dentilis. Movable digit with three teeth. Synarthrodial membrane 
rounded. 

Pedipalps. Paraxial border of trochanter with a slightly thickened ridge, ending 
near pectinate seta v2, seta vl pilose (Fig. 11 G). Anterolateral seta of femur spatulate 
and pectinate on one edge; genu with anterolateral setae simply spatulate. 

Legs. Structure and setation unremarkable. Coxa II provided, on anterolateral 
face, with ridge of fine denticles plus one solitary denticle. Measurements: tarsus I = 
164-168 um; tarsus IV = 173-183 um. 

Discussion. H. ellipticus is very similar in morphology to H. gibber, as well as 
to four known species: A. siculus (Berlese, 1906), H. mallorcae Juvara-Bals, 1975, 
H. maritimus Hyatt, 1987 and H. lawrencei, Hyatt, 1987. These six species likely 
form a monophyletic species-group, which we name mallorcae group with charac- 
teristics listed below. 

The males can be distinguished mainly due to: sternogenital shield without 
excipulum; hypostome (hypostomatic setae on distinct piece of cuticle, hypostome 
more or less evidently extended between corniculi); leg II (spur on femur relatively 
short). 

The females share similar features for: presternal plates (anterior margin serra- 
ted, lateral platelets free); endogynium (cup-shaped, circular, with two prolongations 
from posterior margin); structure of genital shields ( paragynia devoid of thickening in 
front of coxa IV, similar pattern of subapical epigynial structure); structure of chelicera 
(3 and 5 denticles on movable and fixed digits, respectively). 


Differential diagnosis of the Holoparasitus mallorcae species-group. 

Male: 
le corniculi indented, tectum with 3 prongs; mobile digit of chelicera with 4-6 
denticles; fixed digit denticulated: H. mallorcae Juvara-Bals, 1975 
corniculi indented, tectum with | prong; mobile digit of chelicera curved, 
enlarged in its upper third with 6 small denticles; fixed digit slender, truncate 
apically, with 2-3 denticles on inner margin: H. siculus (Berlese, 1905) 


D 


NEW SPECIES OF HOLOPARASITUS 29 


(59) 


corniculi indented, tectum triangular; mobile digit of chelicera with 5 small 

denticles distally and 1 larger proximally; fixed digit slightly curved apically 

with 7-8 denticles : H. maritimus Hyatt, 1987 

4. corniculi conical; mobile digit of chelicera with 6-7 denticles, its dorsal margin 
weakly sclerotized proximally, sclerotized microsclerite near basis of spermato- 
trema; fixed digit slender with one teeth and 7 denticles around pilus dentilis : 
H. ellipticus sp. n. 

I corniculi conical; mobile digit of chelicera with | tooth on internal edge and 
with one large protuberance on external edge of digit; fixed digit slender with 4 
denticles and little tooth around pilus dentilis: H. gibber sp. n. 

6. corniculi conical, mobile digit of chelicera with 1-2 teeth on internal edge;fixed 

digit edentate, finger-like: H. lawrencei Hyatt, 1987 


Female: 

Ik apex of epigynium elongated: A. siculus (Berlese, 1905) 

2. apex of epigynium normal; endogynium without teeth, endogynial margin 
prolongations arcuate: H. maritimus Hyatt, 1987 

3: apex of epigynium normal; endogynium without teeth, endogynial margin 
prolongations straight: H. mallorcae Juvara-Bals, 1975 

4. apex of epigynium normal; endogynium with two teeth, prolongations sinuous: 
H. gibber sp. n. 

> apex of epigynium normal; endogynium with several teeth, prolongations 
sinuous: A. ellipticus sp. n. 

6. apex of epigynium enlarged with little tip medially; endogynium with many 
internal teeth, prolongations curved, usually with denticles or corrugations at the 
end: A. lawrencei Hyatt, 1987 

ACKNOWLEDGEMENTS 


We express our sincere appreciation to the following persons for providing us 
with the material used in the descriptions: Dr. C. Athias-Henriot, Miremont-Haut 
Mauzens, France; Drs M. Castagnoli, F. Pegazzano and R. Nannelli, Experimental 
Institute of Agricultural Zoology, Florence, Italy; Prof. Dr. R. Dallai, University of 
Siena, Siena, Italy; Prof. Dr. V. Mahnert, Dr. B. Hauser, Museum of Natural History, 
Geneva, Switzerland, Dr. L. Tiefenbacher, Munich, Germany. We are also indebted to 
Drs F. Pegazzano and M. Castagnoli for some observations on type material in Berlese 
Acaroteca. For reviewing the manuscript and for their helpful comments, we kindly 
thank Dr. E. Lindquist, Centre for Land and Biological Resources Research, Ottawa, 
Canada and Dr. I. Löbl, Museum of Natural History, Geneva. 


REFERENCES 


BERLESE, A. 1906. Monographia del genere Gamasus Latr. Redia 3: 66-304. 

Evans, G. O. & TILL, W. M. 1979. Mesostigmatic mites of Britain and Ireland (Chelicerata: 
Acari - Parasitiformes). An introduction to their external morphology and classification. 
Transactions of the Zoological Society of London 35: 139-270. 


30 ILINCA JUVARA-BALS & WOJCIECH WITALINSKI 


Hyatr. K. H. 1987. Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: Para- 
sitidae) in the British Isles. Bulletin of the British Museum (Natural History), Zoology 
series 52: 139-164. 

JUVARA-BALS, I. 1975. Sur le genre Holoparasitus Oudemans et sur certains caracteres morpho- 
logiques de la famille Parasitidae Oudem. (Parasitiformes). Acarologia 17: 384-409. 

JUVARA-BALS, I. 1995. Holoparasitus (Holoparasitus) vasilei n. sp. from Israel (Acari, Gamasida: 
Parasitidae) (pp. 47-52). In: Nitzu, E. (ed.) Soil Fauna of Israel. Editura Academiei 
Romäne 47-52. 

LinDauIsT, E. E. & Evans, G. O. 1965. Taxonomic concepts in the Ascidae, with a modified setal 
nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Memoirs of the 
Entomological Society Canada 47: 1-64. 

MICHERDZINSKI, W. 1969. Die Familie Parasitidae Oudemans, 1901 (Acarina,Mesostigmata). 
Panstwowe Wydawnictwo Naukowe, Krakow, 690 pp. 

SCHMÖLZER, K. 1991. Landmilben aus Kärnten I. (Acarina, Parasitiformes). Carinthia II, 181: 
343-358. 

SCHMOLZER, K. 1995a. Einige neue Landmilben aus Südkärnten (Acari,Parasitiformis) Atti del 
Museo Civico di Storia Naturale di Trieste 46: 99-112. 

SCHMÖLZER, K. 1995b. Landmilben aus den östlichen Karawanken (Acarina, Parasitiformis) 
Carinthia II, 185: 499-512. 

Turk, F. A. 1953. A synonymic catalogue of British Acari. Annals and Magazine of Natural 
History 6: 1-26, 81-99. 

VAN DER HAAMMEN, L. 1980. Glossary of Acarological Terms. Vol.I. General Terminology, 
W. Junk Publishers, The Hague, 244 pp. 

VINNIK, O. M. 1994. A new species of the genus Holoparasitus (Mesostigmata, Parasitiformes) 
from the Crimea. Vestnik Zoologii Kiev 2: 82-84. 

VITZTHUM, H. v. 1923. Acarologische Beobachtungen. 7. Reihe. Archiv für Naturgeschichte 
89A: 97-181. 

WITALINSKI, W. 1994a. Holoparasitus (Holoparasitus) dallaii sp. n., a new gamasid mite from 
Sardinia, Italy (Acarı: Pergamasidae). International Journal of Acarology 19: 349-354. 

WITALINSKI, W. 1994b. Two new Holoparasitus species from Italy and Poland (Acari: Gamasida: 
Pergamasidae). Genus 5: 215-222. 


REVUE SUISSE DE ZOOLOGIE 107 (1): 31-48; mars 2000 


Three new species of Geodia Lamarck, 1815 
(Porifera, Demospongiae) 
from the bathyal depths off Brazilian coast, Southwestern Atlantic 


Carla Maria Menegola da SILVA! & Beatriz MOTHES? 

| Universidade de Säo Paulo, Museu de Ciéncias Naturais, Fundaçäo Zoobotänica do 
Rio Grande do Sul, Av. Salvador Franga, 1427, 90690.000, Porto Alegre, RS, Brasil. 
E-mail: silva@ portoweb.com.br 

2 Museu de Ciéncias Naturais, Fundacäo Zoobotanica do Rio Grande do Sul, 
Caixa Postal 1188, 90001-970, Porto Alegre, RS, Brasil. 
E-mail: bmothes @ portoweb.com.br 


Three new species of Geodia Lamarck, 1815 (Porifera, Demospongiae) 
from the bathyal depths off Brazilian coast, Southwestern Atlantic. - 
This work comprises a taxonomic study of tetractinellid sponges from a 
poorly known region in the southwestern Atlantic, off the Rio Grande do 
Sul State coast, Brazil (31°05’—32°00’S/49°31’-50°00’W). Samples were 
collected by R/V “Atlantico Sul” of Fundagäo Universidade do Rio Grande 
in a continental slope survey (“Projeto Talude”) and R/V “Prof. W. Bes- 
nard”, of Instituto Oceanografico da Universidade de Sao Paulo/Group for 
the Development of the Fisheries Industry, in Rio Grande do Sul I Cruise. 
Three new species are described: Geodia australis, G. splendida and 
G. riograndensis. 


Key-words: Porifera - Geodia - Rio Grande do Sul - Brazil - taxonomy - 
continental slope. 


INTRODUCTION 


Six species of Geodia are registered for the Brazilian coast: Geodia gibberosa 
Lamarck, 1815 (Laubenfels, 1956), Geodia neptuni (Sollas, 1886, 1888 as Synops 
neptuni; Mothes, 1996), Geodia tylastra Boury-Esnault, 1973 (Boury-Esnault, 1973), 
Geodia papyracea Hechtel, 1976 (Hechtel, 1976), Geodia corticostylifera Hajdu et al., 
1992 (Hajdu ef al., 1992) and Geodia glariosa (Sollas, 1886, 1888 and Volkmer-Ribeiro 
& Mothes-de-Moraes, 1975 as Cydonium glariosus). 

The present paper describes three new species dredged off Rio Grande do Sul 
State coast (31°05’-32°00°S/49°31’-50°00’W) (fig. 1), in the slope region (207 to 
520 m depth), during oceanographic expeditions carried out by R/V “Atlantico Sul”, of 


Manuscript accepted 12.11.1999 


32 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES 


Fundacäo Universidade do Rio Grande, Projeto Talude; and by R/V “Prof. W. Bes- 
nard”, of Instituto Oceanogräfico, Universidade de Säo Paulo, in agreement with 
Group for the Development of the Fisheries Industry, in Rio Grande do Sul I Cruise. 

The influence of the Subtropical Convergence, with marked seasonal latitu- 
dinal displacement, characterizes the southern/southeastern Brazilian continental shelf 
and slope regions (23°S-34°S) as a biogeographic transition zone (Mothes, 1996; 
Sharp, 1988) between the large neritic areas of Patagonia and tropical Brazil. The 
composition and abundance of species, the pelagic structure, the spatial distribution of 
communities and their trophic interactions, as well as biological production are 
largely controlled by the seasonal dominance of distinct water masses over shelf and 
slope. The studied material was collected in the summer period (october to april), 
when the influence of Tropical Waters is greatest, though waters of subantarctic 
origin may also rise during the summer along the southernmost shelf break regions. 

The paratypes of Geodia australis sp. n. were collected between Sarita and Rio 
Grande localities, 101 Km off Rio Grande do Sul coast, with temperature of 14,50°C 
and salinity 35,76%c. The holotype of this species was collected between Mostardas 
and Solidao localities, 58 Km off Rio Grande do Sul coast. Temperature and salinity 
data for this sample are not known, as well as for the type-specimens of Geodia 
splendida sp.n. and G. riograndensis sp. n. 


MATERIAL AND METHODS 


The samples are deposited in the Porifera Collection of Museu de Ciéncias 
Naturais, Fundaçäo Zoobotanica do Rio Grande do Sul (MCN/ FZB). 
Abbreviations used are: 
BMNH Natural History Museum, London 
MCN/FZB Museu de Ciéncias Naturais, Fundaçäo Zoobotanica do Rio Grande do 
Sul, Porto Alegre, Brazil 
MHNG Muséum d histoire naturelle Genéve, Switzerland 
UFRJ Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil 
ZMA Zoölogisch Museum Amsterdam, Netherlands 


The methodology used to prepare thick sections and dissociated spicules slides 
follow Mothes-de-Moraes (1978). Electron micrographs were taken at MCN/FZB 
with a Jeol 5200 equipment, with an accelerating voltage of 25 kV and magnifications 
varying from 1,500 to 10,000 times. Spicule measurements refer to minimum, mean, 
and maximum sizes in micrometers (um) and were obtained by taking 50 measures of 
each type of spicule/specimen (unless stated otherwise). 


DESCRIPTIONS 
Order Astrophorida 
Geodiidae Gray, 1867 
Geodia Lamarck, 1815 


THREE NEW SPECIES OF GEODIA 33 


40°W 


South America 


Fic. 1 


Map of South American Coast with the Rio Grande do Sul State Coast in detail, showing the 
geographic distribution of Geodia australis sp. n. (MW), G. splendida sp. n. (A) and G. riogran- 
densis sp. n. (D)). 


Type-species: Geodia gibberosa Lamarck, 1815 

Definition: Geodiidae with afferent and efferent aquiferous system indepen- 
dently, with well developed and large subectosomal spaces. Megascleres triaenes. 
Microscleres sterrasters and euasters of different types. Sterrasters varying from 
flattened or globose young forms, smooth with many spherical spaces or provided of 
conical and short rays, with blunt or strongiliform ends sometimes presenting small 
distal holes, to globose forms provided of star or rosette like microspinature at the distal 
end (adapted from Desqueyroux-Fatindez & Van Soest, 1997). 


Geodia australis sp. n. Figs 1, 2, 5, 7, 11-26 


Material: Holotype MCN 330, R/V “Prof. W. Besnard”, off Rio Grande do Sul State 
coast, St. 458 (33°29’S/50°44’ W), 9/XII/1968, 207 m, rocky substrate; Paratype MHNG-INVE 


34 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES 


26564 (schizoparatype-slides deposited in the MCN 331), same data of the holotype; Paratype 
ZMA POR13418 (schizoparatype-slides deposited in the MCN 332), R/V “Prof. W. Besnard”, 
off Rio Grande do Sul State coast, St. 444 (31°31’00”S/49°47’00”W), 6/XII/1968, 284 m, rocky 
substrate. 

Description: Spherical sponge (fig. 2) (diameter 3.8 cm x 3.1 cm, height 2.8 
cm). Hispid surface, with slight brushes of oxeas (fig. 7); small openings are observed 
in some points of the surface, which could not be differentiated in ostia or oscula. 
Colour in spirit grayish-white; compressible consistency. The sponges are associated 
with polychaets and corals. 

Skeleton: Ectosomal (fig. 5): cortex (0.5 mm thick) with spherasters of variable 
sizes, over several layers of sterrasters ended at the cladome-layer of the dichotriaenes, 
plagiotriaenes and protriaenes; scattered small oxeas, strewn at random, are also 
observed. 

Choanosomal (fig. 5): formed by dichotriaenes, plagiotriaenes, protriaenes and 
large oxeas, the last ones preferentially arranged in radial bundles, forming a right angle 
with the cortex. Besides the triaenes and oxeas, sterrasters, spherasters and oxyasters 
are abundantly spread. 


Spicules: Oxeas I (fig. 12): fusiform, straight or slightly curved, with pointed or 
blunt ends; some with mucronate ends; length 1403-2285. 7-3818/24-31.0-40 um. 

Oxeas II (fig. 13): fusiform, with gradually pointed ends; length 181-275. /- 
418/3.2-4.8-6.9 um. 

Dichotriaenes (fig. 11): conical rhabdom, with gradually pointed or strongyloid 
end. Cladi are first curved upwards and then slightly downwards; rhabdome 1012- 
2246.0-3565/33-61.0-86 um: cladome 333-553.8-703 um, protocladi 161-/92./- 
238/29-38.7-48 um; deuterocladi 76-/07.2-143 um. 

Protriaenes (fig. 17): conical, thin rhabdome, with gradually pointed or blunt 
end; cladi with thin ends, sometimes provided with a constriction; rhabdome 1334- 
3143.9-5865/4.6-10.4-23 um; cladome 86-/54.2-276 um, cladi 67-156.4-276/4.6-8.2- 
14 um. 

Plagiotriaenes (fig. 16): rare (N=5), rhabdome conical, straight and thin, with 
gradually pointed end; cladı with thin ends; rhabdome 828-1909/19-28.5 um; cladome 
105-219 um; cladi 51-131/11.5-18.4 um. 

Anatriaenes (fig. 14): rhabdome conical and thin, with pointed or strongyloid 
end; cladi with pointed ends, sometimes marked by a constriction; rhabdome 1150- 
3450.0-6140/4.8-14.2-24 um; cladome 67-/30.9-190 um, cladi 48-//0.6-181 um. 
Some rares (N=5), slender and smaller anatriaenes (fig. 15) can be observed: rhabdome 
© 323-423/3.4-4.6 um; cladome 6.9-9.2 um; cladi 2.3-4.6/1.1-1.8 um. 

Sterrasters (figs. 18-20): oval or more rarely spherical, the young scleres with 
microspined surface in the shape of pointed cones and in the shape of a star in adult 
scleres; diameter 266-3 /4.5-352/190-253.6-295 um. 

Spherasters (fig. 21): spherical with well delimited center, variable size and 
short conical spines with blunt ends; in the choanosome, the largest spherasters can be 
taken for young sterrasters, differing by the shape of the microspines; diameter 20-26.3- 
32 um. 


THREE NEW SPECIES OF GEODIA 35 


FIGs 2-4 


Habit: 2, Geodia australis sp. n.; 3, G. splendida sp. n.; 3, G. riograndensis sp. n.-inner view in 
transversal section. Scale = 2 cm. 


36 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES 


Oxyasters I (fig. 22): small and smooth center; 4 to 7 long, conical and micro- 
spined rays; total diameter 35-43.2-52 um, diameter of the centrum 2.3-4.3-6.9 um, 
rays 9-16.9-25/1.5-2.3-3.5 um. 

Oxyasters II (fig. 23): small and smooth center: 8 to 11 long, conical and micro- 
spined rays; total diameter 16-23.7-32 um, diameter of the centrum 2.3-4.2-5.8 um, 
rays 6.9-/0.0-13.8/2.3 um. 

Spheroxyasters (fig. 26): discrete center; 15 to 22 short , conical and scarcely 
microspined rays; total diameter 14-/6./-20 um, diameter of the centrum 4.6-5.7-6.9 


Ria clear And smooth; 7 to 14 short, blunt rays, with conical 
A total diameter 4.6-7.0-9.2 um, diameter of the centrum 2.3-2.8-3.4 um, 
rays 1.8-2.3-2.8/<1.0 um. 


Etymology: The specific name refers to the type-locality, off Brazil’s southern 
region [Latin Bai australis = southern: south]. 


Remarks: The samples utilized in this description were identified by Mothes-de- 
Moraes (1978) as Geodia eosaster (Sollas, 1886). 

Hajdu et al. (1992) advanced the idea that G. eosaster sensu Mothes-de-Moraes 
(1978) could be a new species due to its widely disjunct distribution when compared to 
the original record of G. eosaster from Australia (Sollas, 1886; 1888). Our reexa- 
mination of Mothes-de-Moraes (1978) specimens, when compared with the syntype of 
G. eosaster [BMNH 1889.1.1.87], revealed several spicular micrometric distinctions 
and the SEM analysis confirmed the presence of adittional categories of megascleres 
and microscleres. We thus described them as Geodia australis sp. n. 

Both species share the presence of dichotriaenes, protriaenes, spherasters and 
strongylasters, but they can be distinguished by the occurrence of rare plagiotriaenes, 
oval shaped sterrasters, and of a second type of oxyaster in the new species. 

From the Tropical western Atlantic records of Geodia, the new species appears 
closest to G. spherastrea Lévi, 1964, from deep-waters off Puerto Rico, at 2840 m 
depth. Both species share the presence of dichotriaenes, protriaenes, anatriaenes, spher- 
asters and strongylasters, but can be distinguished by the occurrence of rare plagio- 
triaenes, oxyasters and oval shaped sterrasters in the new species. 

The “somal” spherasters of Geodia eosaster (Sollas, 1886) and the chiasters 
[=strongylasters sensu Boury-Esnault & Riitzler, 1997] of G. spherastrea (Lévi, 1964), 
correspond to the spherostrongylasters here described for G. australis sp. n. The term 
was coined for cases where the width of the aster’s centrum exceeds 1/3 of the 
microscleres total diameter. 


Geodia splendida sp. n. Figs 1, 3, 6, 8, 27-39 


Material: Holotype MCN 2355 (schizoholotype MHNG-INVE 26565), R/V “Atlantico 
Sul”, off Rio Grande do Sul State coast, St. 10 (32°00°S/50°00’W), 10-X-1991, 520 m, rocky 
bottom. 

Description: Globose sponge (fig. 3), diameter 27 cm, height 24 cm. Surface 
smooth to the touch; single oscule, apical, central, elypsoidal, diameter 3.5/3 cm, at the 


THREE NEW SPECIES OF GEODIA 37 


Serum 2353581 


Fics 5-8 


Skeleton: 5, 6.Skeletal arrangement. 5, Geodia australis sp. n.; 6. G. splendida sp. n.; 7, 8. 
Sponge surface : 7, G. australis sp. n.; 8, G. splendida sp. n. 


38 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES 


159182 


Fics 9-10 


Geodia riograndensis sp. n.: 9, skeletal architecture: 10, oxeas protracting at the sponge surface. 


THREE NEW SPECIES OF GEODIA 39 


Il 


12 8 


Mi 


Fics 11-17 


Megascleres of Geodia australis sp. n.: 11, dichotriaene; 12, oxeas I; 13, oxea II; 14, anatriaene: 
15. smaller anatriaene; 16, plagiotriaene; 17, protriaene. Scales = 100 um. 


—— 


17 


40 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES 


terminal part of a cylindrical channel (length 12 cm), in the inner lateral walls of which 
the exhalant channel openings can be observed; pores not visible. Preserved material of 
violet colour and hard consistency. 


Skeleton: Ectosomal (fig. 6): cortex made up of several overlaping layers of 
sterrasters (0.6-0.7 cm) and the discrete protraction of robust oxeas and some rare 
styloid forms (fig. 8); the cladomes of the orthotriaenes are placed parallel to the 
sponge surface just below the cortex. 

Choanosomal (fig. 6): formed by the rhabdoms of the orthotriaenes, perpendi- 
cular to the surface and, among them, long oxeas, randomly distributed and rare 
sterrasters. 


Spicules: Oxeas I (fig. 27): straight or slightly bent, robust, with ends blunt or 
gradually pointed, length 2254-2681.0-3151/ width 28-44.2-62 um. 

Oxeas II (fig. 30): straight or slightly curved, with gradually pointed or mucro- 
nate ends; some thinner scleres have blunt ends. Length 228-432.7-684 um, width 5.7- 
10.7-19 um. 

Orthotriaenes (figs 28, 29): straigth; cladı straight or bent downward at their 
distal portion, with blunt or gradually sharpening ends; length 3266-3689.0-4094 um, 
width 104-///.6-120 um, cladome diameter 920-//65.7-1495 um.; cladi length 437- 
589.8-759 um, cladi width at the base 85-/00.8-113 um. 

Sterrasters (figs 31-36): spherical or ellipsoidal, with conspicuous hilum, micro- 
spined at the outer portion (figs 34, 35); surface with rounded holes (figs. 31, 32) or 
conical rays (fig. 33) in young forms; or provided with regular microspinature in star 
shape in adult scleres (figs 34-36); diameter 95-148.2-171/86-//9.7-152 um. 

Oxyasters I (fig. 37): 4 to 8 microspined rays all along their length; total 
diameter 78-/00./-131 um, center 6.9-9.2-13.8 um, rays 35-46.9-62/2.8-4.7-6.9 um. 

Oxyasters II (fig. 38): 4 to 9 rays provided with conical microspines all along 
their length, total diameter 12-/7.9-23 um, center 1.6-2.3-3.4 um, rays length 4.6-7.9- 
11.5 um, rays width < | um. 

Spherostrongylasters (fig. 39): 5 to 11 microspined rays all along their length, 
with blunt ends, diameter 5.5-7.4-9.9 um, rays 1.1-/.5-2.2/<1.0-1.5 um. 


Etymology: The specific name refers to the beauty and large size of the 
specimen. [Latin word splendidus = magnific] 


Remarks: Geodia splendida sp. n. is close to Geodia corticostylifera Hajdu et 
al., 1992 [Holotype UFRJ POR 3098 and Paratype UFRJ POR 3714, examined] by the 
shared presence of oxeas, orthotriaenes and oxyasters. They both differ nevertheless by 
the presence of an additional category of styles instead of oxeas in G. corticostylifera 
and of microscleres of the spheroxyaster and strongylospheraster types in G. splendida 
sp. n. 


Geodia riograndensis sp. n. Figs 1, 4, 9, 10, 40-58 


Material: Holotype MCN 1591 (schizoholotype MHNG-INVE 26566),“R/V Atlantico 
Sul”, off Rio Grande do Sul State coast, St. 2-26 (31°05’S/49°31’W), 15-II-1987, 300 m, rocky 
substrate; Paratype MCN 3452, R/V . “Atlantico Sul”, off Rio Grande do Sul State coast, St. 1-5 
(32°24°55”S/50°14°85”W), 30/1V/1986, 200 m, rocky substrate. 


THREE NEW SPECIES OF GEODIA 4] 


25V x1.,000 


Sun 8533214 


— # Se ' 
25kU *1-560 18mm 633221 pus Sue 033228 


€. 


X7r 500 


Fics 18-26 


Microscleres of Geodia australis Sp. n.: 18, young sterraster; 19, adult sterraster surface; 20, adult 
sterraster with hilum; 21, spheraster and spherostrongylaster (arrow); 22, oxyaster I; 23, oxyaster 
II; 24, spherostrongylaster with conical tips; 25, spherostrongylaster; 26, spheroxyaster. 


42 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES 


28 


29 


30 


Fics 27-30 


Megascleres of Geodia splendida sp. n.: 27, oxea I; 28, orthotriaene; 29, orthotriaene cladome in 
apical view; 30, oxeas II. Scale = 500 um. 


THREE NEW SPECIES OF GEODIA 43 


.i1Sku x3.See 


2355514 


Fics 31-39 


Microscleres of Geodia splendida sp. n.: 31, 33 sterraster developmental stage; 32, sterraster 
developmental stage surface; 34, adult sterraster; 35, sterraster surface with hilum; 36, adult 
sterraster showing microspinature details; 37, oxyaster I; 38, oxyaster II: 39, spherostrongylaster 
developmental stage (arrow) and grown spherostrongylaster. 


44 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES 


Description: Subglobose fragment (fig. 4), diameter 10.6 cm, height 7.5 cm. 
Hispid surface, with slight brushes of oxeas (0.2-0.5 mm) (fig. 10); central oscule 
(diameter 11 mm), raising above the surface (3 mm), at the terminal part of a cylindric 
channel (length 44 mm), in the inner lateral wall of which the opening of the exhallant 
channels can be observed; pores were not observed. Preserved material with beige 
colour and firm consistency. 


Skeleton: Ectosomal (fig. 9): Cortex made up by large quantities of sterrasters in 
regular overlapping layers, throughout which, small and thin oxeas are found and, more 
rarely, very long oxeas with filiform ends, both projecting slightly above the sponge 
surface. In the subcortical area orthotriaenes and additional categories of triaenes are 
distributed slant or perpendicular to the surface, the latter being hardly observed. 

Choanosomal (fig. 9): formed by tracts of oxeas perpendicular to the surface 
and rare sterrasters of random distribution. 


Spicules: Oxeas I (fig. 42): thin, sinuous, with gradually pointed ends, length 
2415-5720.5-8464 um, width: 12-/7.5-23 um. 

Oxeas II (fig. 43): robust, straight or slightly curved, with blunt or mucronate 
ends. Some scleres thinner, with one of the ends presenting lateral conical expansions, 
from which one of the sclere sides becomes gradually thinner, length 1610-2082.0-2726 
um, width 21-35.2-46 um. 

Oxeas III (fig. 44): straight or slightly curved, with gradually pointed ends, 
length 247-486.6-741 um, width 5.7-8.9-11.4 um. 

Orthotriaenes (figs 40, 41): straight rhabd with end conical or sharpening gra- 
dually; straight or downwards cladi with conical or strongyliform distal ends, some- 
times bi- or trifurcate, length 1725-28/9.8-3675 um, width 44-66.5-92 um, cladome 
diameter 575-775.3-989 um, cladi length 253-365.4-437 um, cladi thickness at the base 
32-55.0-69 um. 

Anatriaenes (fig. 50): rare (N=6); straight rhabd with gradually pointed ends, 
cladi with gradually pointed or strongyliform ends, length 4501 um, width 9.5-12.3 (m, 
cladome diameter 33-67 um, cladi length 19-38 um, cladi thickness at the base 6.6-9.5 
um. 

Plagiotriaenes (fig. 49): rare (N=4); straight rhabd with gradually pointed end: 
cladi gradually pointed, length 1495-1886 um, width 28-39 um, cladome diameter 460- 
506 um, cladi length: 230-253 um, cladi thickness at the base 23-37 um. 

Protriaenes (figs 47, 48): rare (N=6), straight or slightly curved rhabd, with blunt 
or abruptly pointed end, length 3030-5282 um, width: 9.5-19 um; cladome diameter 
95-204.2 um, cladi length 62-124 um, cladi thickness at the base 8.5-14.2 um. 

Anamesotriaenes (fig. 51). rare (N=5); straight rhabdome with abruptly pointed 
or strongyliform end; cladi with conical or strongyliform ends, length 5938-7581 um, 
width 9.5- 14.2 um, cladome diemeter 49-67 um, cladi length 19-38 um, cladi 
thickness at the base 6.7- 14.2 um. 

Promesotriaenes (fig. 45): rare (N=3); straight rhabdome with gradually pointed 
end; cladi with gradually pointed or strongyliform end, length 2484-3404 um, width 
9.5-19 um, cladome diameter 105-190 um, cladi length 52-105 um, cladi thickness at 
the base 9.5-16.1 um. 


THREE NEW SPECIES OF GEODIA 45 


We 


| 44 
43 if 
47 | 
16 he 
48 
4g i 
i 50 | 
at: I | 51 
Fics 40-51 


Megascleres of Geodia riograndensis sp. n.: 40, orthotriaene; 41, orthotriaene cladome in apical 
view; 42, oxea I; 43, oxeas II; 44, oxeas III; 45, promesotriaene; 46, diaene; 47, protriaene; 48, 
protriaene basal extremity; 49, plagiotriaene; 50, anatriaene; 51, anamesotriaene. Scales = 
200 um. 


46 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES 


25kU x3,See Sem 159107 


*3,588 159182 


[i 
F 
3 


LIALT EL 


25kU x*7.588 ibm 159111 


FIGS 52-58 
Microscleres of Geodia riograndensis sp. n.: 52, sterraster, oxyaster I and oxyaster II (arrow); 53, 
sterraster surface with hilum; 54, oxyaster I and II (arrow); 55, oxyaster II e spherostrongylaster 
(arrow); 56, strongylaster and spherostrongylaster (arrow); 57, strongylaster; 58, strongylaster 
(arrow) and spherostrongylaster. 


THREE NEW SPECIES OF GEODIA 47 


Diaenes (fig. 46) : rare (N=2); straight or slightly sinuous rhabdome, with 
gradually pointed end, length 184 um, width 17 um, cladome diameter 200 um, cladi 
length 157 um, cladi thickness at the base 13.3 um. 

Sterrasters (figs 52, 53): spherical or oval; hilum spherical and conspicuous; 
surface provided with irregular microspines in form of a rosette, sometimes absent in 
the region around the hilum: diameter 57-98. /-124 um/48-89.9-114 um. 

Oxyasters I (figs 52, 54): 3 to 8 microspined rays distributed along its whole 
length, diameter 64-86,4-117 um, center: 5.7-8.3-11.5 um, rays length 30-4/.9-58 um, 
rays width 2.3-3.8-4.6 um. 

Oxyasters II (figs 52, 54, 55): 4 to 9 microspined rays at the distal portion or, 
more rarely, all along their length, diameter 14-20.2-30 um, center 1.8-3.0-4.6 um, rays 
length 4.6-9.0-13.8 um, rays width 1.1-/.6-2.3 um. 

Strongylasters (figs 56-58) - varying to spherostrongylasters (figs 55, 56, 58). 5 
to 13 rays with strongyliform or truncate ends, microspined all along their length or, 
more rarely, at the distal half, diameter 4.6-8.4-13.2 um, center 1.6-3.2-4.6 um, rays 
length: <1.0-3.0-4.6/< 1.0 um. 


Etymology: The specific name refers to the Rio Grande do Sul State coast, off 
which the sponges were collected. 


Remarks: When compared with other species of Geodia from the Brazilian 
coast, Geodia riograndensis sp. n. is close to G. glariosa Sollas, 1886 [Syntype BMNH 
1889.1.1.86] by the shared occurrence of oxeas, ortho-, pro- and anatrienes as mega- 
scleres, differing, however, for presenting additional categories of megascleres, as three 
categories of oxeas, plagiotriaenes, anamesotriaenes, promesodiaenes, promesotriaenes 
and diaenes; and of microscleres, as oxyasters, strongylasters and strongylospherasters. 


ACKNOWLEDGEMENTS 


We are thankful to Fundaçäo de Amparo a Pesquisa do Estado do Rio Grande do 
Sul (FAPERGS) for granting the Master’s Newly-Graduated Scholarship to the senior 
author (Proc. n° 96/60364-1) in the first part of the work (study of Geodia riogran- 
densis sp. n. and G. splendida sp. n.) and Aid to the junior author (Proc. n° 95/ 0728.3); 
to Fundaçäo de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP) for granting the 
Doctoral’s Scolarship (study of Geodia australis sp. n. and conclusion of the paper) to 
the senior author (Proc. n° 98/00797-4); to Dr. Ricardo R. Capitoli (FURG), for the 
donation of the specimens studied; to Dr. Clare Valentine (BMNH) and Dr. Guilherme 
Muricy (UFRJ), for lending the type-materials intended for comparative studies; to the 
Museum of natural history of Geneva (MHNG) for cooperation; to Dr. Eduardo Hajdu 
(MNRJ) and M. Sc. Cléa B. Lerner (USP), for their critical reading and suggestions for 
the paper; to the technicians Cleodir J. Mansan and Marcia Spadoni (MCN), for the 
scanning electron microscope photos and to Mrs. Rejane Rosa (MCN) for the final 
scleres drawings and map. 


48 CARLA MARIA MENEGOLA DA SILVA & BEATRIZ MOTHES 


REFERENCES 


BOURY-ESNAULT, N. 1973. Campagne de la Calypso au large des côtes atlantiques de 1° Amérique 
du Sud (1961-1962). I, 29. Spongiaires. Results of scientific Campaign of Calypso 10: 
263-295. 

Boury-ESNAULT, N. & RÜTZLER, K. 1997. Thesaurus of Sponge Morphology. Smithsonian 
Contributions to Zoology 596: 1-55. 

DESQUEYROUX-FAUNDEZ, R. & VAN SOEST, R. W. M. 1997. Shallow waters Demosponges of the 
Galäpagos Islands. Revue suisse de Zoologie 104 (2): 379-467. 

HaJpu, E. C. M, Muricy, G., CusToDIo, M., Russo, C. & PEIXINHO, S. 1992. Geodia 
corticostylifera (Demospongiae, Porifera) New Astrophorid from the brazilian coast 
(Southwestern Atlantic). Bulletin of Marine Science 51 (2): 204-217. 

HECHTEL, G. J. 1976. Zoogeography of Brazilian Marine Demospongiae (pp. 237-259). In: 
Harrison, F. W. & Cowden, R. R. (eds). Aspects of Sponge Biology. 

LAUBENFELS, M. W. DE. 1956. Preliminary discussion of the sponges of Brazil. Boletim do 
Instituto Oceanografico 1: 1-4. 

Levi, C. 1964. Spongiaires des zones bathyale, abyssale et hadale (pp. 63-112). In: Torben Wolff 
(ed.). Galathea Report, Scientific results of the Danish Deep-Sea Expedition Round the 
World, 1950-1952. 

MOTHES, B. 1996. Esponjas da Plataforma Continental Norte e Nordeste do Brasil. Unpublished 
thesis, Universidade de Sao Paulo, Instituto de Biociéncias, Sao Paulo, SP, 230 pp. 

MOTHES-DE-MORAES, B. 1978. Esponjas tetraxonidas do litoral sul-brasileiro: II. Material cole- 
tado pelo N/Oc. “Prof. W. Besnard” durante o Programa RS. Boletim do Instituto 
Oceanografico 27 (2): 57-78. 

SHARP, G. D. 1988. Fish populations and fisheries (pp. 155-202). In: H. Postma & J.J. Zijlstra 
(eds). Continental Shelves. Ecosystems of the World, vol. 27. Elsevier, Amsterdam. 

SOLLAS, W. J. 1886. Preliminary account of the tetractinellid sponges dredged by H.M.S. 
“Challenger”, 1872-76. Part I. The Choristida. Scientific Proceedings of the Royal Dublin 
Society 5 (4): 177-199. 

SOLLAS, W. J. 1888. Report on the Tetractinellida collected by the H. M. S. “Challenger” 
during the years 1873-1876. Report on the scientific results of the voyage of H. M. S. 
“Challenger” Zool. 25: xi-clxvi + 1-457 pp. 

VOLKMER-RIBEIRO, C. & Mothes-de-Moraes, B. 1975. Esponjas tetraxonidas do litoral sul-brasi- 
leiro. I - Redescricào de Cydonium glariosus Sollas, 1886 e Erylus formosus Sollas, 
1886. Iheringia, ser. Zool. (47): 3-22. 


REVUE SUISSE DE ZOOLOGIE 107 (1): 49-79; mars 2000 


Oribatids from Switzerland III (Acari: Oribatida: Oppiidae 1 
and Quadroppiidae). 
(Acarologica Genavensia XCIII) 


Sandor MAHUNKA & Luise MAHUNKA-Papp 
Zoological Department, Hungarian Natural History Museum, Baross utca 13, 
H-1088 Budapest, Hungary. 


Oribatids from Switzerland III (Acari: Oribatida: Oppiidae 1 and 
Quadroppiidae). (Acarologica Genavensia XCHI). — Oppioid oribatids 
taken from soil samples in Switzerland are recorded. There are 15 species, 
of which 6 are new for science and one of them also represents a new 
genus: Paramedioppia gen. n. (Oppiidae). The following 5 new combina- 
tions are proposed: Berniniella conjuncta (Strentzke) comb. n. et stat. n. = 
Oppia sigma conjuncta Strentzke, 1951; Berniniella hauseri (Mahunka) 
comb. n. = Oppia hauseri Mahunka, 1974; Berniniella serratirostris 
(Golosova) comb. n. = Oppia serratirostris Golosova, 1970; Dissorhina 
signata (Schwalbe) comb. n. = Oppia signata Schwalbe, 1989; Lauroppia 
maritima (Willmann) comb. n. = Oppia maritima Willmann, 1929; 1 spe- 
cies is placed in synonymy: Oppiella rafalskii Optotna & Rajski, 1983 = 
Berniniella hauseri (Mahunka, 1974): syn. n. In addition 2 species (still not 
recorded for Switzerland) are discussed, in this way a total of 17 oppioid 
species are considered. Morphological and distributional data of 11 species 
are provided and the nature of relationships and some additional morpholo- 
gical characters are discussed. 


Key-words: Acari - Oribatida - Oppiidae - Quadroppiidae - taxonomy - 
new species - new genus - new combinations - Switzerland. 


INTRODUCTION 


Our revision work regarding the oribatids of Switzerland, which will eventually 


be incorporated into a book, has several times been mentioned (e.g. Mahunka, 1993, 
1996a). For the very simple reason that at least another 4-5 years will have to elapse 
until the appearence of this book, we believe that all the taxonomic novelties and 
specific faunistic data should be published in order to complement the ever increasing 
number of taxonomic and zoogeographic researches. This time we propose to discuss 
a part of the available information gained in connection with species belonging to the 
superfamily of Oppioidea. 


Manuscript accepted 13.01.1999 


50 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


We should stress that, in referring to the taxon superfamily, we adopt the 
system incorporating e.g. the following families: Oppiidae, Quadroppiidae and 
Suctobelbidae. In this paper we discuss especies belonging to the families Oppiidae 
Grandjean, 1951 and Quadroppiidae Balogh, 1983. The subfamilial division within 
the family Oppiidae seems quite unsuitable for further subdivisions, especially since 
there are many genera (see Mahunka, 1999). 


List of localities 


GE-4 = Switzerland: Genève: Frontenex, pieds souches chênes; 14.VII.1980; leg. C. Besuchet 
— (33). 

GR-6 = Switzerland: Graubünden: Landquart, pieds aulne (Alnus); 26.1X.1983; leg. C. Besu- 
chet — (118). 

GR-8 = Switzerland: Graubiinden: Samnaun, alpine Wiesen mit Rhododendron, Gesiebe, 
2050m; 26.VIIL. 1968; leg. C. Besuchet — (26). 

GR-9 = Switzerland: Graubünden: Santa Maria — Paß Umbrail, Gesiebe, 2000m; 5.VIII.1974; 
leg. C. Besuchet — (37). 

GR-10= Switzerland: Graubünden: Untervaz b. Chur, mousses: 29.IX.1983; leg C. Besuchet — 
(126). 

LU-I = Switzerland: Luzern: Eigenthal, pres Eigenthal (village) Forenmoos, Sphagnum, 
970m.; 2.VIIL. 1996; leg. C. Besuchet — (108). 

NW-1 = Switzerland: Nidwald: Musenalp, oberhalb Niederrickenbach, mousses dans lappiaz, 
1800m; leg. C. Besuchet — (113). 

NW-2 = Switzerland: Nidwald: Pilatus, tamisage rhododendron, 1800m; 14.VI.1984; leg. 

I. Löbl — (123). 

Switzerland: Nidwald: Trübsee, mousses près source, 1800m; 8.1X.1997; leg. 

C. Besuchet — (127). 

SO-5 = Switzerland: Solothurn: Schottwill, Bucheggberg, Rindenmoos von lebendem Baum; 
27.1X.1987; leg. S. Mahunka & L. Mahunka-Papp — (47). 

SZ-3 = Switzerland: Schwyz: Pragelpaß, mousses sapins, 1650m; 25.VIII.1983; leg. I. Lobl — 
(116). 

TI-5 = Switzerland: Tessin: Monadello - Moneto, im faulenden Laub, 850m; 23.IV.1992; 
leg. C. Besuchet — (91). 

TI-9 = Switzerland: Tessin: Nufenen-Paß, Boden, dürres Laub und Baummulm aus einem 
Lärchenwald; 15.VI.1979: leg. S. Mahunka & L. Mahunka-Papp - (18). 

TI-11 = Switzerland: Tessin: Rancate, forêt de chätaigniers, tamisages; 7.1X.1965; leg. 
C. Besuchet — (25). 

VS-4 = Switzerland: Valais: Daubensee, mousses et herbes, 2200m; 11.VIII.1980; leg. 
C. Besuchet — (32). 

VS-8 = Switzerland: Valais: Forêt de Finges, souches pins (Pinus); 14.VIII.1980; leg. 
C. Besuchet — (5). 


NW-3 


EIST OF IDENTIFIED SPECIES 


OPPIIDAE Grandjean, 1951 


Berniniella conjuncta (Strentzke, 1951) comb. et stat. n. 
Locality: NW-2. 
Distribution: Germany: first record for Switzerland. 
Berniniella hauseri (Mahunka, 1974) comb. n. 
Localities: GR-6; TI-11. 
Distribution: Europe: first record for Switzerland. 


ORIBATIDS FROM SWITZERLAND III 5] 


Dissorhina signata (Schwalbe, 1989) comb. n. 
Locality: GR-9. 
Distribution: Germany (known from the type locality only); first record for 
Switzerland. 
Lauroppia hauseri sp. n. 
Localities: TI-5; TI-11. 
Lauroppia maritima (Willmann, 1929) comb. n. 
Localities: SO-5; TI-9. 
Distribution: Palearctic Region; first record for Switzerland. 
Lauroppia obscura sp. n. 
Localities: GR-8; VS-4; VS-8. 
Moritzoppia incisa sp. n. 
Localiy: NW-3. 
Moritzoppia keilbachi (Moritz, 1969) 
Locality: SZ-3. 
Distribution: Europe; first record for Switzerland. 
Oppiella besucheti sp. n. 
Locality: SZ-3. 
Oppiella propinqua sp. n. 
Locality: LU-1. 
Oxyoppioides decipiens (Paoli, 1908) 
Locality: GE-4. 
Distribution: Central and Southern Europe, Caucasus; first record for Switzerland. 
Paramedioppia helvetica gen. n., sp. n. 
Locality: TI-9. 
Subiasella quadrimaculata (Evans, 1952) 
Locality: GR-10. 
Distribution: Europe; first record for Switzerland. 


QUADROPPIIDAE Balogh, 1983 


Quadroppia longisetosa Minguez, Ruiz & Subias, 1985 
Locality: NW-1. 
Distribution: Southern Europe; first record for Switzerland. 
Quadroppia michaeli Mahunka, 1977 
Locality: not recorded for Switzerland. 
Distribution: Greece and Spain. 
Quadroppia omodeoi Mahunka & Paoletti, 1984 
Locality: not recorded for Switzerland. 
Distribution: Italy. 
Quadroppia cf. paolii Woas, 1986 
Locality: GR-10. 
Distribution: Palearctic Region; first record for Switzerland. 


DESCRIPTION AND REDESCRIPTION OF SOME OF THE OPPIOID SPECIES 


The present paper records 15 species belonging to the superfamily Oppioidea 
Grandjean, 1951 found in the territory of Switzerland (13 species of the family Oppii- 
dae and 4 species of the family Quadroppiidae). In exceptional cases we also discuss 
species unrecorded in Switzerland so far, when it is deemed important to elucidate 
certain specific taxonomic questions. Among the species 6 are new for science and | 
represents a new genus (Paramedioppia). In addition some extremely rare species are 
touched upon. Morphological complementations, corrections, new combinations, the 
publication of synonyms and the reinterpretation of some species are also given. 


52 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Berniniella conjuncta (Strenzke, 1951) comb. et stat. n. Figs 1-3 


Oppia sigma conjuncta Strenzke, 1951: 723, fig. 5. 
Oppiella sigma (Strentzke, 1951): sensu Woas, 1986: 182, Abb. 88-89. 

Material examined: Switzerland: NW-2. 

Remarks: The recently examined soil samples harboured, in many cases, 
the very same species which we could readily identify with the description of 
Strenzke’s Oppia sigma conjuncta published in 1951, and with Woas's redescription 


Fics 1-3 


Berniniella conjuncta (Strenzke, 1951) — 1: body in dorsal view, 2: body in ventral view, 3: 
podosoma in lateral view. 


ORIBATIDS FROM SWITZERLAND III 53 


of Oppia sigma published in 1986. Without doubt Strenzke's description is inadequate 
since the description of the ventral side is missing. However, on the basis of the Swiss 
specimens and the redescription of Woas now we may safely conclude that O. 
conjuncta is an independent, valid species. Furthermore, we should like to point out 
that Woas made a mistake when preparing his redescription, since his specimens 
belong to O. conjuncta and not to O. sigma. 

The drawings were made from the specimens originating from Switzerland 
which readily correspond with the figures of both Strenzke and Woas. For this reason 
a complete redescription would be out of place here. Nevertheless, we should like to 
draw the attention to some important features. 

Measurements: Length of body: 208-224 um, width of body: 102- 
114 um. 

Dorsal side (Fig. 1): Rostrum with wide median apex, lateral apices 
much smaller. Costulae S-shaped, directed to the bothridia. One pair of median laths 
directed outwards, not connecting with the costulae. Bothridium with posterior 
tubercle, behind it comparatively large pustules are evident. Sensilli with short 
bristles. Dorsosejugal suture protruding anteriorly, ten pairs of notogastral setae 
nearly equal in length. 

Lateral part of podosoma (Fig. 3): Well sclerotized, longitudinal 
crests and pustulated or granulated fields are visible. Ratio of prodorsal setae: ro = ex 
> in = le. Setae /c do not arise on pedotecta I, discidium well developed bearing 
setae 4c. 

Ventral side (Fig. 2): Sternal apodemes and borders weakly developed 
or absent. Sejugal apodemes with longitudinal, arched lines in opposite position. 
Posterior margin of epimeral borders 2 and 4 undulate or denticulate. Epimeres 1 
framed laterally by longitudinal crests bearing setae /c anteriorly. Anogenital setal 
formula: 4 - 1 - 2 - 3. All setae in the epimeral and anogenital region simple, setiform, 
without conspicuous cilia. 


Berniniella hauseri (Mahunka, 1974) comb. n. Figs 4-5 


Oppia hauseri Mahunka, 1974: 585, Abb. 34-36. 
Oppiella rafalskii Optotna & Rajski, 1983: 543, figs 1-8. Syn. n. 


Material examined: Switzerland: GR-6; T-11. 

Remarks: The recently collected specimens are quite identical with those 
described from Greece (Mahunka, 1974). However, the ventral side was not 
described, so the relegation of the species was only provisional owing to lack of 
information regarding the number of genital setae and some other ventral features. 
After having again examined the type species we found, as it was expected from the 
dorsal side, that O. hauseri has 4 pairs of genital setae corresponding well in all other 
features with those of specimens collected recently in Switzerland. One of these was 
selected for the drawings (Figs 4-5). 

Taxonomic position: After examining the related species, it is 
certain that Berniniella serratirostris (Golosova, 1970) comb. n. and Berniniella 


54 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


RE 


\ ) 
Fics 4-5 


Berniniella hauseri (Mahunka, 1974) — 4: body ın dorsal view, 5: body in ventral view. 


rafalskii (Optotna & Rajski, 1983) comb. n. are its closest allies. Peculiarly enough, 
we could not find any differentiating character between the latter and B. hauseri, thus, 
we consider them synonyms; so the latter is a junior synonym of B. hauseri. The 
authors’ description - besides the obvious mistakes - 1s wholly identical with the 
features of the Swiss specimens. The only difference we could ascertain stems from 
the drawing technique, due to some simplified method used by the authors. Thus, the 
median part of the dorsosejugal region, the costula on the basal part of the prodorsum 
and also the median thickening are figured to be more complicated than they in fact 
are. A similar problem is found with the drawing of the sejugal borders and the 
coxisternal region. The position of setae 2b, 2c and 3c is obviously erroneously 
depicted. Optotna & Rajski (1983) thoroughly evaluated the relationship of Oppia 
serratirostris with Oppiella rafalskii. These differences (especially the number of 
branches of the sensilli, the shape of the costulae and the shape of the median rostral 
apex) are convincing enough for us, too. 


Dissorhina signata (Schwalbe, 1989) comb. n. Figs 6-9 


Oppia signata Schwalbe, 1989: 99, Abb. 1. 
Material examined: Switzerland: GR-9. 


ORIBATIDS FROM SWITZERLAND III 


Nn 
Un 


Fics 6-7 
Dissorhina signata (Schwalbe, 1989) — 6: body in dorsal view, 7: body in ventral view. 


Remarks: On the basis of the sensillar shape and its unilateral ciliation, 
the species of the genus Dissorhina Hull, 1916 may grouped into two. The “ornata 
species group” has bacilliform sensilli, whose distal end is spiculate or quite smooth, 
while the “fricarinatoides species group” has sensilli with very long unilaterally 
arranged branches. The studied species belongs to the second group. 

Redescription: The specimens of this species collected in Switzerland 
have been stored for some time in the Geneva collection. These specimens are easily 
identifiable with the description and the given figures. Some important features which 
deserve special attention are described below. 

Measurements. — Length of body: 197-208 um, width of body: 103- 
134 um. 

Dorsal side (Fig. 6): Rostrum tripartite, rostral setae arising exactly on 
the triangular, well separated median apex. Costulae casket-shaped, diverging me- 
dially and converging distally. Lamellar setae arising on small separated parts. Bothri- 
dium with a posterior tubercle, behind it a well developed, separated, round tubercle. 
Sensilli with 6-7 long, branches unilaterally. Dorsosejugal suture well protruding 
anteriorly, with a straight median part. Ten pairs of notogastral setae nearly equal in 
length. Setae c, not shorter than the others. 


56 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Fics 8-9 
Dissorhina signata (Schwalbe, 1989) — 8: podosoma in lateral view, 9: leg I. 


Lateral part of podosoma (Fig. 8): Well sclerotized, longitudinal 
crests and pustulated or granulated fields are visible. Ratio of prodorsal setae: ro = ex 
>in = le. Setae /c do not arise on pedotecta I, discidium well developed and bearing 
setae 4c. 

Ventral side (Fig. 7): Sternal apodemes absent between bo. 2 and bo. 
sej. Sejugal apodemes with longitudinal, arched lines. Posterior margin of epimeral 
borders 4 undulate or denticulate. Epimeres 1 framed laterally by longitudinal crests 
bearing setae /c. Anogenital setal formula: 5 - 1 - 2 - 3. All setae in the epimeral and 
anogenital region simple, setiform, without conspicuous cilia. 

Legs: Form and chaetotaxy of leg I as shown in Fig. 9. 

Taxonomic position: The species D. signata stands very near to D. 
tricarinatoides (Dubinina, in Dubinina er al. 1966) (see also Mahunka, 1996b). It may 
be distinguished by the shape of the prodorsal costulae with the auxiliary comple- 
mentary ribs, which are present in D. tricarinatoides and absent in D. signata, and the 
position of setae ad, which arise nearer to the anal plates than the lateral margin of 
ventral plate in D. signata (much farther, near to the lateral margin in D. trticari- 
natoides). 


ORIBATIDS FROM SWITZERLAND III 


57 


Fics 10-12. Lauroppia hauseri sp. n. — 10: body in dorsal view, 11: body in ventral view, 
12: podosoma in lateral view. 


Lauroppia hauseri sp. n. Figs 10-12 


Material examined: Switzerland: Holotype: Tessin: TI-11, 1 paratype from 


the same sample; 6 paratypes: Tessin: TI-5. Holotype and 4 paratypes: MHNG!, 3 paratypes 
(1614-PO-98): HNHM?. 


' MHNG: deposited in the Muséum d’histoire naturelle, Geneva. 


2 HNHM: deposited in the Hungarian Natural History Museum, Budapest, with identification 
number of the specimens in the Collection of Arachnida. 


58 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Diagnosis: Rostrum elongated, rostral apex roundish. Short, basally 
dilated, guttiform costulae present Sensilli pectinate. Median part of the anterior 
notogastral margin straight. Ten pair of notogastral setae, setae c, long. Six pairs of 
genital setae, anogenital region normal. 

Measurements: Length of body: 369-443 um, width of body: 188- 
214 um. 

Prodorsum: Rostral part of prodorsum conspicuously elongated, rostral 
apex conical, rounded medially. Prodorsal surface with short basal transcostulae pro- 
truding from the guttiform basal tubercles. Bothridia well developed, with conspi- 
cuously large basal tubercles. Rostral setae arising from the rostral surface, lamellar 
setae located very far from the rostrals, near to the interlamellar setae. Lamellar setae 
fine, setiform, arising at the distal end of the costulae, interlamellar setae much 
thicker, ciliate, straight, arising from the guttiform tubercles. Sensilli pectinate, direc- 
ted forwards, each bearing 7 branches unilaterally. 

Lateral part of podosoma: Sejugal region covered by a thick 
cerotegument layer. Surface weakly granulated, with larger pustules between the 
acetabula. Acetabula IV placed above acetabula I-III. Setae /c arising near to small 
pedotecta I (Fig. 12). 

Notogaster: Anteromedian margin of notogaster straight, not penetrating 
between the bothridia. A weak crista present. Ten pairs of notogastral setae present, 
setae c, long, longer than the others. Setae /a arising in front of Im (Fig. 10). Setae la 
and h; longer than the remaining ones, setae p,, p,, and p3 shortest of all. 

Ventral side (Fig. 11): Apodemes and borders weakly developed, 
epimeral surface ornamented by a weak polygonal design. Posterior border of epi- 
meral region undulate or denticulate. Epimeral setae partly ciliate or bifurcate. Lateral 
setae (3c, 4c) longer than the inner ones. Anogenital setal formula: 6 - 1 - 2 - 3. All 
setae in anogenital region simple and short. 

Remarks: On the basis of the prodorsal structure and the basal position of 
the lamellar and interlamellar setae the new species belongs to the “fallax-group” (see 
Willmann, 1931). The new species is distinguished from its congeners by the ratio 
and form of the prodorsal setae and the very short / and p setae. 

Derivatio nominis: We dedicate the new species to our friend, Dr. B. 
Hauser, the initiator of the “Oribatida Helvetica project”. 


Lauroppia maritima (Willmann, 1929) comb. n. Fig. 13-15 


Oppia maritima Willmann, 1929: 45, Abb. 4. 
Oppia maritima: Strenzke, 1951: 720, fig. 2 

Material examined: Switzerland: SO-5; TI-9. 

Remarks: The species was described by Willmann (1929), and Strenzke 
(1951) redescribed it. Both these descriptions are inadequate and some very important 
features are missing. Therefore, we give herewith some drawings and a short des- 
cription. 


ORIBATIDS FROM SWITZERLAND III 59 


Fics 13-15 


Lauroppia maritima (Willmann, 1929) — 13: prodorsum in dorsal view, 14: epimeral region, 
15: podosoma in lateral view. 


Prodorsum: Rostral apex sharply pointed, the incisure wide, lateral 
apices much smaller than the median one (Fig. 13). Exobothridial setae bifurcate. 
Exobothridial region well sclerotized (Fig. 15), surface around the bothridia and 
acetabulae I-IV pustulate or granulate. 

Notogaster: Protruding anteriorly, however, the median part of the 
dorsosejugal line is straight. 

Ventral side: Posterior border of the coxisternal region with pustulate 
margin, pustules are in a transversal hollow on both sides (Fig. 14). Setae /c arising 
far from pedotecta I. Setae 3c bifurcate. Anogenital setal formula: 5 - 1 - 2 - 3. 
Anterior pair of genital setae much longer than the others. 


60 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Lauroppia obscura sp. n. Figs 16-20 


Material examined: Switzerland: Holotype: Valais: VS-8, 6 paratypes from 
the same sample; 4 paratypes: Valais: VS-4; 6 paratypes: Graubünden: GR-8. Holotype and 10 
paratypes: MHNG, 6 paratypes (1615-PO-98): HNHM. 

Diagnosis: Rostrum divided by two incisions, median apex triangular. 
Prodorsum with complicate costulae. Sensilli clavate, unilaterally spinose. Notogaster 
straight anteriorly. Ten pairs of notogastral setae, c, long. 5 pairs of genital setae, 
anogenital region normal. 

Measurements: Length of body: 280-305 um, width of body: 
148-172 um. 

Prodorsum: Rostral apex (Fig. 17) divided by two small incisions, 
median apex conspicuously wide, sometimes triangular, or blunt at tip, sometimes its 
margin undulate. Lateral apices with simple, sharp points. Among the costulae one 
pair arched and comparatively well developed, they do not reach the bothridium or 
the bases of the lamellar setae. A pair of S-shaped basal costulae also present 
reaching to the inner margin of the longitudinal ones (Fig. 20). Lateral part of 
prodorsum well sclerotized, with some tubercles and pustules, one pair of which bears 
the exobothridial setae. In the dorsosejugal region one pair of indistinct porose areas 
also visible (Fig. 19). Bothridia large with strong posterior apophyse. Rostral setae 
arising dorsally, comparatively near to each other. Lamellar setae shorter than the 
other prodorsal setae, arising on small tubercles independently of the longitudinal 
costula. Sensilli elongate, their head asymmetrically clavate, with short spines on their 
outer margin. 

Lateral part of podosoma: Exobothridial region and fields 
between acetabula I - IV granulated by comparatively large granules or pustules. 
Some well-sclerotised, long, longitudinal crests also present. A pair of porose areas in 
the sejugal region. Positions of acetabula III and IV nearly the same as those of ace- 
tabula I and II (Fig. 19). Pedotecta I small, setae /c arising far from its basal part. 
Discidium well developed, setae 4c arising far from its lateral margin, on the epimeral 
surface. 

Notogaster: Elongated, dorsosejugal part straight anteriorly. Crista 
clearly seen. Ten pairs of notogastral setae present, setae c, conspicuously long, 
setiform, all others somewhat shorter. Setae /m arising in front of la (Fig. 16). Setae 
pP], P and pz only slightly shorter than the others. 

Ventral side (Fig. 18): Epimeral region hardly sclerotised. Apodemes 
partly absent only borders visible between the borders 2 and sejugal borders or sejugal 
borders and borders 4. Lateral borders on epimeres | arched bearing setae /c. Sejugal 
borders broad, with a pair of hollows medially, with similar, but smaller ones, also 
present on bo 2. All epimeres with a polygonal pattern. Setae 4c arising on the 
epimeral surface. Epimeral setae long, simple. Anogenital setal formula: 5 - 1 - 2 - 3. 
Among the genital setae the two anterior pairs conspicuously long, directed forwards. 
Position and shape of the aggenital, anal setae and lyrifissures iad normal. 

Legs: Solenidia of tibia IV very long and curved . 


ORIBATIDS FROM SWITZERLAND III 61 


Fics 16-20 


Lauroppia obscura sp. n. — 16: body in dorsal view, 17: rostrum, 18: body in ventral view, 
19: podosoma in lateral view, 20: basal part of prodorsum. 


62 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Remarks: On the basis of the form and structure of the prodorsum the new 
species belongs to the L. maritima group. Although closely related to L. maritima, the 
head of the sensillus of the new species is much longer and the spines on the sensillar 
head are very short (long in L. maritima). 

Derivatio nominis: Named after the complicated prodorsal design 
and rostral apex. 


Moritzoppia incisa sp. n. Figs 21-23 


Materialexamined: Switzerland: Holotype: Nidwald: NW-3, 26 paratypes 
from the same sample. Holotype and 16 paratypes: MHNG, 10 paratypes (1613-PO-98): 
NHMH. 


Diagnosis: Rostrum roundish. Prodorsum with complicate costulae. 
Sensilli clavate, unilaterally pectinate. Anterior margin of notogaster protruding into 
the interbothridial region and interrupted medially. Cristae present. Ten pairs of 
notogastral setae, setae c, long. Four pairs of genital setae, anogenital region normal. 

Measurements: Length of body: 267-276 um, width of body: 152- 
155 um. 

Prodorsum: Rostrum slightly elongated, with nasiform median apex. 
Lamellar costulae weakly sclerotized, lamellar apices usually separated, their anterior 
part carrying some pustules. A pair of elongated, guttiform tubercles in the inter- 
bothridial region bearing interlamellar setae at their outer margin. A pair of round 
tubercles present in the middle part of the costulae, they are opposed on both sides 
(Fig. 21). Some weak sigillae visible laterally. Bothridia well developed, with large 
basal tubercles surrounded with a granulated field. Rostral setae arising on the dorsal 
surface, ciliate. Lamellar setae arising on the lamellar apices. The ratio of the pro- 
dorsal setae: ro > in = ex > le. Sensilli with well developed heads, bearing 9-10 
branches unilaterally. 

Lateral part of podosoma: Well sclerotized, nearly the whole 
surface well granulated and/or pustulate. Large pustules present also above the ace- 
tabula of legs HI and IV (Fig. 23). Some longitudinal crests also present. Ventral 
border especially undulate and incised, acetabula I-IV arranged nearly in a longitu- 
dinal line. No porose area visible behind the bothridia. 

Notogaster: Dorsosejugal part of the notogaster protruding anteriorly in 
the interbothridial region. Crista present laterally. Dorsosejugal margin interrupted 
medially and framed by a pair of short crests directed posteriorly, sometimes some 
spots are visible in this slit. Ten pairs of notogastral setae present, setae c, slightly 
longer than the others, la, Im and /p equal in length, setae h and setae p only slightly 
shorter. 

Ventral side (Fig. 22): Well sclerotized, all apodemes and borders well 
developed. Lateral margin of epimeres | framed by longitudinal borders, but setae /c 
arising on pedotecta I. The surface of epimeres I pustulate, posterior borders also 
conspicuously covered by granules or pustules. Sejugal borders with some pairs of 
longitudinal crests or lines. Posterior margin of borders IV strongly undulate. Setae 4b 
arising at the posterolateral corner. Surface around discidium also pustulate. Setae 4c 


ORIBATIDS FROM SWITZERLAND III 63 


Fics 21-23 


Moritzoppia incisa sp. n. — 21: body in dorsal view, 22: body in ventral view, 23: podosoma in 
lateral view. 


64 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


stand further from the discidia. All epimeral setae simple and smooth. Anogenital 
setal formula: 4 - 1 - 2 - 3. All setae simple. Lyrifissures iad short, located in paraanal 
position. 

Remarks: On the basis of the structure of the dorsosejugal margin of the 
notogaster, the new species is distinguished from all congeners. 

Derivatio nominis: Named after the incised anterior margin of the 
notogaster. 


Moritzoppia keilbachi (Moritz, 1969) 


Oppia keilbachi Moritz, 1969: 37, Abb. 1-3. 
Moritziella keilbachi: Balogh, 1983: 26, 9: 9p-v. 
Moritzoppia keilbachi: Subias & Rodriguez, 1988: 122. 


Material examined: Switzerland: SZ-3. 


Remarks: The species was described by Moritz (1969) on the basis of 
females. A male specimen has been found in the Swiss material. The habitus of the 
male is more slender and it is also smaller (210um x 92 um) than the female. In all 
other features it corresponds well with the original description and the figures. Espe- 
cially important is the granulate surface behind the bothridium and on the discidium 
and the shape of the sensillus. The sensillus of the studied individual is blunt at tip, its 
head bearing simple, short cilia, arranged in 2-3 rows. Features of the ventral region 
also accord well with the original description, e.g. the posterior borders of the 
coxisternal region have an undulate margin. 


Oppiella besucheti sp. n. Figs 24-27 


Material examined: Switzerland: Holotype: Schwyz: SZ-3, 5 paratypes from 
the same sample. Holotype and 3 paraypes: MHNG, 2 paratypes (1611-PO-98): NHMH. 


Diagnosis: Rostral apex divided by two incisions, median apex 
triangular. Prodorsum with costulae and other structures. Sensilli narrow, fusiform 
with lateral branches. Anterior margin of notogaster straight, with lateral protube- 
rances. Ten pairs of notogastral setae, setae c, long. Five pairs of genital setae, 
anogenital region normal. 

Measurements: Length of body: 250-259 um, width of body: 99- 
105 um. 

Prodorsum: Rostrum divided by two small, but wide, U-shaped 
incisions, median apex triangular, also small (Fig. 24). Median costulae Y-shaped, 
weakly developed. Behind them, in the interbothridial position, one pair of elongated, 
nearly guttiform tubercles. Along the lamellar costulae, at their outer side, a pair of 
opposed smaller tubercles present on each side. Bothridia with posterior tubercles. 
The whole surface of prodorsum distinctly granulated or pustulated, also in the 
lamellar region (Fig. 27). All four pairs of prodorsal setae long, interlamellar ones 
thicker and more heavily ciliated than the others. Ratio among them: in > ro = ex > 
le. Sensilli very long, slightly dilated medially, bearing 5-6 lateral branches of 
different lengths on their distal parts. 


ORIBATIDS FROM SWITZERLAND III 65 


Fics 24-27 


Oppiella besucheti sp. n. — 24: body in dorsal view, 25: body in ventral view, 26: podosoma in 
lateral view, 27: basal part of prodorsum. 


66 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Lateral part of podosoma: Well sclerotized, nearly the whole 
surface well granulated and/or pustulated. Some longitudinal crests also present. 
Ventral border especially waved and incised, acetabula I-IV arranged nearly in one 
longitudinal line (Fig. 26). A pair of porose areae behind the bothridia clearly seen. 

Notogaster (Fig. 24): With one pair of granulate humeral apophyses. 
Dorsosejugal suture straight. Ten pairs of notogastral setae present, setae c, shorter 
than Ja, Im and Ip also shorter than c,, but nearly equal in length with the other setae, 
excepting p, and p3. Setae h, characteristically directed outwards. Setae c, with cilia, 
all the other setae smooth. 

Ventral side (Fig. 25): Well sclerotized, all apodemes and borders - 
excepting a part of the sternal ones - between the 2nd and sejugal apodemes well deve- 
loped. Lateral margin of epimeres | framed by longitudinal borders bearing setae /c. A 
pair of inner costulae, parallel with discidium also present on epimeres 4 continued as 
undulate transversal parts. Setae 4b arising on their posterolateral corner. Among the 
epimeral setae, setae /c, 3c, but primarily and firstly setae 4c, with long cilia. All other 
epimeral setae simple and smooth. Anogenital setal formula: 5 - 1 - 2 - 3. All setae 
simple. Lyrifissures iad long, placed in paraanal position. 

Legs: Setae of trochanters of legs II well arched, with conspicuously long 
cilia. Setae p of all tarsi simple, setiform. Solenidium @ of leg IV, very long, arched, 
longer than the joint. 

Remarks: See after the following species. 

Derivatio nominis: I dedicate the new species to Dr. C. Besuchet 
(Muséum d'histoire naturelle, Geneva), the collector of the specimens and most of 
those soil samples from Switzerland which we studied recently in Geneva. 


Oppiella propinqua sp. n. Figs 28-32 

Material examined: Switzerland: Holotype: Luzern: LU-1 15 paratypes from 
the same sample. Holotype and 10 paratypes: MHNG, 5 paratypes (1612-PO-98): NHMH. 

Diagnosis: Rostrum nasiform. Lamellar costulae directed to the 
bothridia, one pair of basal protuberances present. Sensilli slightly fusiform, narrow, 
bearing short spines. Anterior margin of the notogaster straight, with well developed 
lateral protuberances. Ten pairs of notogastral setae, setae c, long. Anogenital region 
typical for the genus. Five pairs of genital setae. 

Measurements: Length of body: 263-270 um, width of body: 140- 
145 um. 

Prodorsum: Rostrum slightly elongated, with a nasiform median apex. 
Lamellar costula well sclerotized, covered by pustules on their anterior part. A much 
finer transversal costula between them (sometimes it is hardly observable) (Fig. 30). 
A pair of elongated, guttiform tubercles in the interbothridial region bearing inter- 
lamellar setae on their outer margin and a pair of short laths at an angle between 
costulae and bothridia. Some weak sigillae visible laterally. Bothridia well developed, 
with a basal tubercle with a granulated field around it. Rostral setae characteristically 
arched inwards, ciliate. Lamellar setae arising on the distal half of the costulae. The 


ORIBATIDS FROM SWITZERLAND III 67 


Fics 28-29 
Oppiella propinqua sp. n. — 28: body in dorsal view, 29: body in ventral view. 


ratio of the prodorsal setae: ro > in = ex > le. Sensilli very long, their heads slightly 
dilated medially bearing some (5-7) very short spines. 

Lateral part of podosoma (Fig. 32): Well sclerotized, nearly the 
whole surface well granulated and/or pustulated. Some longitudinal crests also 
present. Ventral border, especially, undulate and incised acetabula I-IV arranged in 
one longitudinal line. A pair of indistinct porose areas behind the bothridia. 

Notogaster (Fig. 28): With one pair of large humeral apophyses. 
Dorsosejugal suture straight medially, slightly arched laterally, near the humeral 
tubercles. Ten pairs of notogastral setae present, setae c, and /a equal in length, /m 
and /p much shorter than the preceding ones. Setae h, characteristically directed 
outwards, no essential difference among setae h and setae p,. Setae c, with short cilia, 
all other setae smooth. 

Ventral side (Fig. 29): Well sclerotized, all apodemes and borders - 
excepting a part of the sternal ones - between the 2nd and sejugal apodemes well 
developed. Lateral margin of epimeres | framed by longitudinal borders bearing 


68 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Fics 30-32 


Oppiella propinqua sp. n. — 30: basal part of prodorsum, 31: epimeral region, 32: podosoma in 
lateral view. 


setae /c. A pair of inner costulae, parallel with the discidium also present on epi- 
meres 4 continued as undulate transversal parts (Fig. 31). Setae 4b arising on their 
posterolateral corner. All epimeral setae simple and smooth. Anogenital setal formula: 
5 - | - 2-3. All setae simple. Lyrifissures iad long, located in paraanal position. 

Legs: Setae p of legs II - IV thick, clearly spiniform. Solenidium of tibia IV 
not longer than the joint. 

Remarks: Oppiella nova (Oudemans, 1902) is one of the most variable 
species of the genus Oppiella Jacot, 1937, however, it was designated as the type of 
the genus. Most authors consider this species as cosmopolitan, but nobody has made a 
comparative study on world material. Our opinion is, that this species comprises a 
number of closely related species. This opinion is strengthened in that some authors 
consider Oppiella uliginosa (Willmann, 1919) as a synonym of O. nova (e.g. Subias 
& Balogh, 1989). On the other hand Woas (1986) in his redescription of O. uliginosa 
quite obviously represents another species. The description and the drawings (Woas, 
1986: 208, Abb. 102-103) surely do not refer to O. nova. 


ORIBATIDS FROM SWITZERLAND III 69 


The two preceding new species stand very near to the type species of this 
genus, and e.g. Oppiella primorica (Golosova, 1970), which also belongs to this 
group. I also regard this genus as rather heterogeneous needing further study to 
demarcate its generic limits. Obviously, on the basis of the O. nova type, the diag- 
nosis of the genus Oppiella should be complemented and restricted so that only those 
species which display the following characteristics should be included: Dorsosejugal 
margin with lateral apophyses. Among the epimeres, epimeres | framed laterally with 
a longitudinal border bearing setae /c, epimeres 4 framed posteriorly by an undulate 
(denticulate) lath, continuing anteriorly as a longitudinal inner costula. Other features 
should be investigated in the future. 

Among the new species, on the basis of the undivided rostrum, Oppiella pro- 
pinqua sp. n. stands closer to Oppiella nova. It is distinguishable from the type spe- 
cies by the form of the sensillus, which is longer in the new species. The other 
differential characters are: form of the prodorsal costula, the form and ratio of pro- 
dorsal setae, length and ciliation of epimeral setae (/c, 3c and 4c). The other species 
(Oppiella besucheti sp. n.) stands nearest to Oppiella primorica (Golosova, 1970) on 
the basis of the divided rostral apex (see also Mahunka, 1979). The new species is 
distinguishable from Oppiella primorica by the much larger median prodorsal apex 
and the thicker and ciliated interlamellar setae and the much wider basal guttiform 
tubercles in the interbothridial region in the new species. 

Derivatio nominis: The new species belongs to the Oppiella nova 
species group. 


Oxyoppioides decipiens (Paoli, 1908) Figs 33-37 
Oppia decipiens Paoli, 1908: 69, figs 29, 48. 

Oppia decipiens: Pérez-Inigo, 1971: 297, fig. 30. 

Oxyoppioides decipiens: Subias & Minguez, 1985: 182. 

Material examined: Switzerland: GE-4. 

Remarks: The species has been collected several times in Central Europe, 
although it is more common in the Mediterraenean area. Several authors (e.g. Pérez- 
Inigo, 1965; Subias & Minguez, 1985) discussed it but no detailed redescription has 
been made. Furthermore,there are no available figures illustrating its leg structure. 
The following remarks complement our knowledge of this species. 

Prodorsum: Rostrum U-shaped, excavated with two small, lateral teeth 
(Fig. 33). Costulae weak, surface densely granulate. Interbothridial region with 
numerous indistinct sigilla. 

Pate ral part of pod os om a. (Eis. 35): Weakly, sclerotized: 
exobothridial region with fine granulation, with only poorly developed, longitudinal 
laths. Exobothridial setae short. 

Notogaster: Humeral processes in the dorsosejugal region, hitherto 
characteristic for the genus Oxyoppia, entirely missing. Anterior margin of notogaster 
arcuate, closed in the middle. 

Ventral side (Fig. 34): Apodemes very poorly developed. Epimeral 
borders hardly discernible in places. From among the sejugal and apodemes 4 and 


70 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Fics 33-35 


Oxyoppioides decipiens (Paoli, 1908) — 33: body in dorsal view, 34: body in ventral view, 35: 
podosoma in lateral view. 


borders the sternal apodeme and border are entirely missing. Surface of discidium 
granulate. Setae /c arising on pedotecta 1. Epimeral setae ciliate, setae 4a and 4b 
located strikingly close to each other. Anogenital formula 5-1-2-3. Aggenital setae 
arising one behind the other. Lyrifissure iad in inverse apoanal position, far removed 
from anal opening. 


ORIBATIDS FROM SWITZERLAND III il 


Legs: Normal, oppioid type, only the tibia of leg I slightly thickened and 
the distal part of femur triangular (Fig. 36), solenidium of tibia of leg IV (Fig. 37) 
conspicuously short, and all setae characteristically long. 

Taxonomic position: Subias & Balogh (1989) placed the genus in 
relationship with Oxyoppia. We do not agree with this opinion, although the genus is 
soundly based. The key published by them is unsuitable for its identification. The 
position of lyrifissure fad, said to be paraanal, is also in error in the key of Balogh & 
Balogh (1992). 


FIGS 36-37 
Oxyopioides decipiens (Paoli, 1908) — 36: leg I, 37: femur, genu and tibia of leg IV. 


Paramedioppia gen. n. 


Diagnosis: Family Oppidae. Rostrum divided by two incisions. Weak 
costulae with a transcostula and one pair of interbothridial “tubercles” present on 
prodorsum. Sensilli setiform, with some irregular spines. Dorsosejugal part of noto- 
gaster gradually arched, not penetrating into the interbothridial region. Crista present, 
setae c, arising medially. Ten pairs of notogastral setae present. Among the epimeral 


Tp. SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


setae /c arising on the epimeral surface, epimeral borders and apodeme normally 
developed. Anogenital setal formula 6 - 1 - 2 - 3. Lyrifissures iad in paraanal position. 
Gnathosoma, chelicerae and palps normal. Legs very long, joints of leg IV especially 
elongated. 

Type species: Paramedioppia helvetica sp. n. 

Remarks: The new taxon seems to be a mixture of two genera: Oppiella 
Jacot, 1937 and Medioppia Subias & Minguez, 1985. The latter is a highly hetero- 
geneous genus, as is Oppiella. The type species of the genus Medioppia, O. media 
Mihelcic, 1956 (Pérez-Iñigo, 1965) does not have costulae but has interbothridial 
tubercles. On the other hand, it has 5 pairs of genital setae, and its rostrum is 
undivided. Thus, we could not relegate the new species into any of the known genera. 

Derivatio nominis: Similar to the genus Medioppia. 


Paramedioppia helvetica sp. n. Figs 38-42 


Materialexamined: Switzerland: Holotype: Tessin: TI-9, 1 paratype from 
same sample. Holotype: MHNG, paratype (1616-PO-98): NHMH. 

Diagnosis: Rostrum divided, rostral apex truncate. Prodorsum weakly 
sclerotized, short longitudinal and transversal costulae present. Sensilli setiform with 
irregular spines. Anterior margin of notogaster convex. Cristae present. Ten pairs of 
notogastral setae, setae c, long. Anogenital setae normal, six pairs of genital setae. 

Measurements: Length of body: 762-823 um, width of body: 369- 
403 um. 

Prodorsum: Elongated. Rostral part rounded, but flanked by two short 
incisions. Median apex truncate, lateral apices rounded. Prodorsal surface weakly 
sclerotized, but short longitudinal costulae with distinct corners, a very weak trans- 
costula and a pair of interbothridial, arched tubercles present. Some indistinct sigilla 
also observable. Bothridia without posterior tubercles. Rostral setae arising on the 
prodorsal surface, lamellar setae located behind the costular corners (Fig. 38). Ratio 
of prodorsal setae: : in = ro > le > ex. Sensilli well characteristic for the species, 
bent several times, setiform, with 4-5 spines, without capitula, their distal ends simple 
or bifurcate. 

Lateral part of podosoma: Pedotecta I comparatively well 
developed. Exobothridial surface not granulate, some granules observable betweeen 
the acetabula II - HI. This part well framed by a lath, above the acetabula (Fig. 40). 

Notogaster: Conspicuously elongated. Crista weak, but distinct also a 
pair of short and weak median lines present at the dorsosejugal margin of the noto- 
gaster (Figs. 38, 40). Ten pairs of notogastral setae of equal length present, all ciliate. 

Ventral side (Fig. 39): Apodemes and borders weakly developed. 
Marginal, longitudinal lath absent on epimera 1. In anteromedian sternal apodemes a 
ring-like feature present on the sternal apodemes, between setae /a. All epimeral 
setae simple, thin, setiform. Epimeral surface ornamented by polygonal pattern. Six 
pairs of genital setae arising in two rows (4 pairs in median, 2 pairs in lateral 
position). Adanal setae ciliate, like the notogastral ones, other setae in the anogenital 
region simple. 


ORIBATIDS FROM SWITZERLAND III 73 


Fics 38-40 


Paramedioppia helvetica gen. n., sp. n. — 38: body in dorsal view, 39: body in ventral view, 
40: podosoma in lateral view. 


Legs: All joints long, not widened, setae conspicuously ciliate/spinose. Leg 
setal formulae are normal for the family: 


I: 1-5-2+1-4+2-20+2 - 1 (Fig. 41) 
IV:1-2-2-3+1-10-1 (Fig. 42) 


74 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Fics 41-42 


Paramedioppia helvetica gen. n., sp. n. 41: leg I, 42: leg IV. 


Remarks: The main features are mentioned in the generic diagnosis, but it 
is the form of the sensillus which well characterises the new species. 

Derivatio nominis: Named after the country the species originates 
from. 


Subiasella (Lalmoppia) quadrimaculata (Evans, 1952) 


n 


Oppia quadrimaculata Evans, 1952: 37, fig. 2. 
Subiasella (Lalmoppia) quadrimaculata: Subias & Rodriguez, 1986: 114, figs 1-3. 

Materialexamined: Switzerland: GR-10. 

Remarks: The species is known from several localities (from England to 
Poland) of Europe. but it is not frequent anywhere. This is its first record from 
Switzerland. The prodorsum of these specimens, as it is the case in the Hungarian 
ones, has neither costula nor lamellar line. 


ORIBATIDS FROM SWITZERLAND III 75 


Quadroppiidae Balogh, 1983 


There is a great confusion in the family Quadroppiidae partly due to the 
publication of Minguez et al. (1985) and partly to Woas (1986) but specifically to that 
of the latter author who synonymized species without the study of types. The core of 
the problem is in the interpretation of the senior author's species (Quadroppia 
michaeli Mahunka, 1977) described from Greece, and on the other hand, Woas' inter- 
pretation of the type species, ©. quadricarinata (Michael, 18853) which was misun- 
derstood by Paoli (1908). Accordingly, if the ©. michaeli species is not new, but 
simply the misinterpreted ©. quadricarinata, then the name introduced by Woas as 
Q. paolii nom. n. is redundant. 

The situation is further aggravated in that the senior author described with 
Paoletti (Mahunka & Paoletti, 1984) another new species belonging to this genus (Q. 
omodeoi Mahunka & Paoletti, 1984), without taking the trouble to point out its 
relationship with the ©. quadricarinata by then correctly understood. Our recent 
study has revealed that Q. paolii is probably identical with the specimens from 
Switzerland, and we further believe that it is not identical either with ©. quadri- 
carinata sensu Mahunka, nor with the form interpreted by Paoli, nor Q. omodeoi. 
There is a further problem in that Minguez er al. (1985) hold the view that Hammer’s 
Q. monstruosa is the same of Paoli’s Q. quadricarinta. 

To clarify the problem thorough study of types is needed along with authenti- 
cally identified specimens. This time we could only examine the types of Q. michaeli 
and Q. omodeoi, and those Greek specimens which were published in 1977 as 
O. quadricarinata sensu Mahunka. We discuss O. longisetosa Minguez, Ruiz et 
Subias, 1985 that was also found in Switzerland (Q. quadricarinata (Michael, 1885) 
was also collected at several localities in Switzerland) but refrain from discussing the 
other species here. 


Quadroppia longisetosa Minguez, Ruiz & Subias, 1985 Figs 43-44 
Quadroppia longisetosa Minguez et al., 1985: 104, figs 6-7. 

Material examined: Switzerland: NW-1. 

Remarks: This species is readily identifiable among the members of the 
genus Quadroppia Jacot, 1939. The Swiss specimens correspond well with the des- 
cription and the figures of the species. Some slight differences exist, like the weak 
longitudinal lath in the interlamellar region parallel with the costulae, and the 
notogastral setae are somewhat more rigid than depicted in the drawing. We provide 
some figures obtained from specimens taken in Switzerland . 


Quadroppia michaeli Mahunka, 1977 Fig. 45 


Quadroppia michaeli Mahunka, 1977: 914, Abb. 12. 
Quadroppia michaeli: Minguez et al., 1985: 114, figs 17-18. 
Quadroppia michaeli: Mahunka, 1977 sensu Woas, 1986: 215. 


> The original description appeared in 1885 and not in 1887 as mentioned by Woas (1986). 


76 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


SSA 
D Sal 


A 
| Tape 
Fics 43-44 


Quadroppia longisetosa Minguez, Ruiz & Subias, 1985 — 43: body in dorsal view, 44: body in 
ventral view. 


Remarks: In spite of Woas’ opinion, this is a valid, independent species. It 
cannot be brought into close relationship with any of the so far described species. Its 
characteristics are: 

Prodorsum: Oval crest in the rostral region undivided and gradually 
becoming narrower basally. On both sides an arcuate, well separated crest present. 
Intercostular region with a short, transverse lath directly behind the distal end of 
costulae. 

Notogaster: The posteriorly running ribs emanating from the humeral 
processes are not weaker than the ones beside them, length of ribs is the same. 

Ventral side: Epimeral borders are shown in Fig. 45. The median 
pattern directly in front of the genital opening is highly characteristic, it comprises 
two touching parts, both are rounded. 

Legs: Tarsus of leg II bears 2 solenidia. 


Quadroppia omodeoi Mahunka & Paoletti, 1984 Figs 46-47 
Quadroppia omodeoi Mahunka & Paoletti, 1984: 114, figs 1-2. 


ORIBATIDS FROM SWITZERLAND III 77) 


Remarks: All the known species are distant relatives only. Its charac- 
teristics are: 

Prodorsum: Oval crest in the rostral region is undivided, not strongly 
narrowing basally. Intercostular region with a short, concave, transverse lath directly 
behind the distal end of costulae. Behind this transverse lath 2 pairs of sigillae and 
one pair of lateral laths are present, the latter are not connected in the middle. Sensilli 
conspicuously long (Fig. 47), much longer than in the other species. 

Notogaster: The posteriorly running ribs emanating from the humeral 
processes are much longer than the lateral ones. Notogaster completely enframed by 
arıb. 

Ventral side: Fig. 46 depicts the epimeral borders. 

Legs: Tarsus of leg II bears 2 solenidia. 


Quadroppia cf. paolii Woas, 1986 Fig. 48-49 
Quadroppia paolii Woas, 1986: 78, figs 30-32. 


Material examined: Switzerland: GR-10. 

Remarks: Without studying the type the relegation of this species is 
impossible. It is highly probable that this species may be identical with Q. monstruosa 
Hammer, 1979 sensu Minguez er al. (1985). Fundamental differences are obvious: 

Prodorsum: Oval crest in the rostral region is either entirely divided 
basally or much variegated. Intercostular region with a long, robust, arcuate lath 
directly before the end of costulae. Just behind this 2 pairs of sigillae and one pair of 
lateral laths, the latter frequently connected in the middle with a weak, transverse lath. 
Sensillar capitula oval (Fig. 49). 


Fics 45-49 


Quadroppia michaeli Mahunka, 1977 — 45: epimeral region. 
Quadroppia omodeoi Mahunka & Paoletti, 1983 — 46: epimeral region, 47: sensillus. 
Quadroppia cf. paolii Woas, 1986 — 48: epimeral region, 49: sensillus. 


78 SANDOR MAHUNKA & LUISE MAHUNKA-PAPP 


Notogaster: The posteriorly running ribs emanating from the humeral 
processes are shorter than the lateral ones. 

Ventral side: as shown in Fig. 48. 

Legs: Tarsus of Leg II bearing 2 solenidia. 


ACKNOWLEDGEMENTS 


First and foremost we should like to thank the collector of this interesting 
material, Dr. C. Besuchet. Our hearthy thanks are due to Dr. V. Mahnert and Dr. B. 
Hauser for the opportunity offered to study the material. We should also like to thank 
Dr. C. Lienhard for the good advice extended on several taxonomic questions. For 
reading the manuscript, translating some parts and linguistically revising others we 
thank Dr. L. Zombori. For the conscientious corrections and the good advice incor- 
porated in the text we extend our sincere thanks to Dr. M. Luxton (National Museum 
of Wales, Cardiff). 


REFERENCES 


BALOGH, J. 1983. A partial revision of the Oppiidae Grandjean, 1954 (Acari: Oribatei). Acta 
Zoologica Academiae Scientarum Hungaricae 29: 1-79. 

BALOGH, J. & BALOGH, P. 1992. The Oribatid mites genera of the world. Hungarian Natural 
History Museum, Budapest, I: 263 pp., II: 375 pp. 

DUBININA, E. V., SOSNINA, E. F., VYSOCKAJA, S. O., MARKOV, G. N. & ATANASOV, L. H. 1966. 
Oribatea aus Nagetiernestern im VitoSa-Gebirge. /zvestiya na Zoologicheskiya Institut 
22: 81-141. 

Evans, G. O. 1952. Terrestrial Acari new to Britain— I. Annals and Magazine of Natural 
History 5: 33-41. 

GoLosova, L. D. 1970. New species of Oribatids (Acariformes, Oribatei) from the South 
Primorye and the Kuril Islands. Zoologichesky Zhurnal 49: 694-701. 

HAMMER, M. 1979. Investigations on the Oribatid Fauna of Java. Biologiske Skrifter. Kongelige 
Danske Videnskabernes Selskab 22: 1-79. 

MAHUNKA, S. 1974. Neue und interessante Milben aus dem Genfer Museum. XII. Beitrag zur 
Kenntnis der Oribatiden-Fauna Griechenlands (Acari). Revue Suisse de Zoologie 81: 
569-590. 

MAHUNKA, S. 1977. Neue und interessante Milben aus dem Genfer Museum XXX. Weitere 
Beiträge zur Kenntnis der Oribatiden-Fauna Griechenlands (Acari: Oribatida). Revue 
Suisse de Zoologie 84: 905-916. 

MAHUNKA, S. 1979. Complementary data to the knowledge of some Oribatid species (Acari). 
Folia Entomologica Hungarica 32: 139-152. 

MAHUNKA, S. 1993. Oribatids from Switzerland I. (Acari: Oribatida). (Acarologica Genavensia 
LXXXD. Archives des Sciences 46: 51-56. 

MAHUNKA, S. 1996a. Oribatids from Switzerland II. (Acari: Oribatida) (Acarologica Gena- 
vensia XC). Folia Entomologica Hungarica 57: 125-129. 

MAHUNKA, S. 1996b. Oribatids of the Bükk National Park (Acari: Oribatida) (pp. 491-532). In: 
Mahunka, S. (ed.). The Fauna of the Bükk National Park. Hungarian Natural History 
Museum, Budapest, vol. II: 655 pp. 

MAHUNKA, S. 1999. Oribatid mites (Acari: Oribatida) from Uganda, II. Arcoppia with 
comments on generic concepts. Acta Zoologica Academiae Scientiarum Hungaricae 45: 
247-272. 


ORIBATIDS FROM SWITZERLAND III 79 


MAHUNKA, S. & PAOLETTI, M. G. 1984. Oribatid mites and other mites (Tarsonemidae, 
Anoetidae, Acaridae) from woods and farms monocultivated with corn in the low 
laying plan (Veneto and Friuli, N-E Italy). Redia 67: 93-128. 

MICHAEL, A. D. 1885. New British Oribatidae. Journal of the Royal Microscopical Society 5: 
385-397. 

MiNGUEZ, M. E., Ruiz, E. & SuBias, L. S. 1985. El género Quadroppia Jacot, 1939, (Acari, 
Oribatida, Oppiidae). Boletin de la Asociation Espanola de Entomologia 9: 95-118. 

Moritz, M. 1969. Neue Oribatiden ((Acari) aus Deutschland V. Oppia keilbachi nov. spec. 
Wissenschaftliche Zeitschrift der Ernst Moritz-Arndt-Universität Greifswald, Mathema- 
tisch-Naturwissenschaftliche Reihe 18: 37-40. 

OPLOTNA, H. & RAISKI, A. 1983. Oppiella rafalskii sp. n. (Acarida, Oribatida). Acta Zoologica 
Cracoviensia 26: 543-561. 

OUDEMANS, A. C. 1902. Acarologische Aanteekeningen. Entomologische Berichten |: 36-39. 

PAOLI, G. 1908. Monografia del genere Dameosoma Berl. e generi affini. Redia 5: 31-91. 

PEREZ-INIGO, C. 1965. Especies espanolas del genero Oppia C. L. Koch (Acari, Oribatei). 
Boletin de la Real Sociedad Espanola de Historia Natural 62 (1964): 385-416. 

PÉREZ-INIGO, C. 1971. Acaros oribätidos de suelos de Espana peninsular e Islas Baleares 
(Acari, Oribatei). Eos. Revista Espanola de Entomologia 46: 263-350. 

SCHWALBE, TH. 1989. Oppiella signata, eine neue Art der Familie Oppiidae aus dem Osterz- 
gebirge (Acari, Oribatei). Deutsche Entomologische Zeitschrift 36: 99-101. 

STRENTZKE, K. 1951. Some new Central European Moss-Mites (Acarina: Oribatei). Annals and 
Magazine of Natural History 4: 719-726. 

SUBIAS, L. S. & BALOGH, P. 1989. Identification keys to the genera of Oppiidae Grandjean, 
1951 (Acari: Oribatei). Acta Zoologica Hungarica 35: 355-412. 

SUBIAS, L. S., MINGUEZ, M. E. 1985. Los Oppidos [sic!] (Acari, Oribatida) de El Pardo (Espana 
central). Serratoppia n. gen. y Oxyoppioides n. gen. Actas do II Congresso Iberico de 
Entomologia (= Suplemento | ao Boletim da Sociedade Portuguesa de Entomologia): 
165-174. 

SUBIAS, L.-S. & RODRIGUEZ, P. 1986. Oppiidae (Acari, Oribatida) de los sabinares (Juniperus 
thurifera) de Espana VI. Neotrichoppia (Confinoppia) n. subg. y Moritziella Balogh, 
1983. Redia 69: 115-130. 

SUBIAS, L. S. & RODRIGUEZ, P. 1988. Oppiidae (Acari, Oribatida) de los sabinares (Janiperus 
[sic!] thurifera) de Espana, VIII. Medioppiinae Subias y Minguez. Boletin de la 
Asociation Espanola de Entomologia 12: 27-43. 

WILLMANN, C. 1919. Diagnosen einiger neuer Oribatiden aus der Umgebung Bremens. 
Abhandlungen herausgegeben vom Naturwissenschaftlichen Verein zu Bremen 24 
(1920): 552-554. 

WILLMANN, C. 1929. Neue Oribatiden II. Zoologischer Anzeiger 80: 43-46. 

WILLMANN, C. 1931. Moosmilben oder Oribatiden (Oribatei). Tierwelt Deutschlands und der 
angrenzenden Meeresteile 22: 79-200. 

Woas, S. 1986. Beitrag zur Revision der Oppioidea sensu Balogh, 1972 (Acari, Oribatei). 
Andrias 5: 21-224. 


7 Le 
x B IE wa ary 4 
u uw Ze ©) é 


1 


an iW “i ii : 
> Abate. Pal 7 
a ER 


Si 
+ WR > 
} CPS 


REVUE SUISSE DE ZOOLOGIE 107 (1): 81-95; mars 2000 


Rudolphiella szidati sp. n. (Proteocephalidea: Monticelliidae, 
Rudolphiellinae) parasite of Luciopimelodus pati (Valenciennes, 
1840) (Pisces: Pimelodidae) from Argentina with new observations 
on Rudolphiella lobosa (Riggenbach, 1895) 


Alicia A. GIL DE PERTIERRA! & Alain de CHAMBRIER? 

! Departamento de Ciencias Biolögicas, Lab. 52, Facultad de Ciencias Exactas y 
Naturales, Pab. II, Ciudad Universitaria, U.B.A., 1428 - Buenos Aires, Argentina. 
e-mail: helminto @bg.fcen.uba.ar 

2 Departement des Invertébrés, Muséum d'histoire naturelle, PO Box 6434, CH -1211 
Geneva 6, Switzerland. 
e-mail: alain.dechambrier @ mhn. ville-ge.ch 


Rudolphiella szidati sp. n. (Proteocephalidea: Monticelliidae, Rudol- 
phiellinae) parasite of Luciopimelodus pati (Valenciennes, 1840) (Pisces: 
Pimelodidae) from Argentina with new observations on Rudolphiella 
lobosa (Riggenbach, 1895). - Rudolphiella szidati sp. n. (Cestoda, Monti- 
celliidae, Rudolphiellinae) is described from the anterior intestine of the 
pimelodid fish, Luciopimelodus pati caught in Parana river and de la Plata 
river (Argentina). Additional informations to the description of the type 
material of Rudolphiella lobosa (type species of the genus) is given. R. 
lobosa was not found in L. pati (as reported originally by Riggenbach, 1895) 
and has never been found since its original description. The new species 
differs from other species of Rudolphiella by its larger scolex, a higher 
number of testes and by its distinct egg shape. A diagnosis of Rudolphiella is 
given. The following taxonomical actions are introduced: Rudolphiella pira- 
catinga (Woodland, 1935) comb. nov. for Monticellia piracatinga Woodland, 
1935; Monticellia rugata Rego, 1975 syn. nov. of Rudolphiella piracatinga 
(Woodland, 1935). The presence of glandular cells within the scolex’s apex 
is observed in all known Rudolphiella species 1.e. R. lobosa, R. myoides, R. 
szidati, R. piracatinga, R. piranabu. Rudolphiella, the sole genus within 
Rudolphiellinae, is widespread in a monophyletic group of hosts (Calophy- 
sus-group sensu de Pinna, 1998) suggesting likely coevolution. 


Key-words: Proteocephalidea — Monticelliidae — Rudolphiella szidati sp. n. 
— Rudolphiella lobosa — Luciopimelodus pati — coevolution — Argentina. 


Manuscript accepted le 9.11.1999 


82 ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER 


INTRODUCTION 


Riggenbach (1895) described Corallobothrium lobosum parasite of Luciopi- 
melodus pati (Valenciennes, 1840) from Paraguay River, Paraguay. Later, Fuhrmann 
(1916) redescribed Riggenbach’s material adding information of proglottides’ 
transverse sections, and erected a new genus Rudolphiella Fuhrmann, 1916 to allocate 
Corallobothrium lobosum Riggenbach, 1895. This species has never been found since 
its original description. 

Woodland (1934) erected the genus Amphilaphorchis and described two species 
Amphilaphorchis piranabu Woodland, 1934 and A. myoides Woodland, 1934, both 
parasites of Pinirampus pirinampu (Spix, 1829). A year later Woodland (1935), 
concluded that genus Amphilaphorchis Woodland, 1934 is a synonym of Rudolphiella 
and erected a new subfamily, the Rudolphiellinae. 

The examination of Luciopimelodus pati (Valenciennes, 1840) from Parana river 
and de la Plata river, Argentina, the type host of R. lobosa (Riggenbach, 1895) revealed 
the presence of a new species of Rudolphiella described herein. Since the original 
description of Rudolphiella lobosa (Riggenbach, 1895) did not contain data on some 
useful morphological structures, the type material of this species 1s restudied. 


MATERIAL AND METHODS 


Sixty-two specimens of Luciopimelodus pati (Valenciennes, 1840) were 
examined for helminths. Forty intestines were placed in lukewarm water in order to 
relax the worms and then fixed in AFA, while others were dissected and fixed directly 
in hot 4% formaldehyde solution and were subsequently stored in ethanol 75 % V/V. 
Entire tapeworms were stained with Langeron’s alcoholic chlorhydric carmine 
(Langeron, 1949), differenciated in acid ethanol, dehydrated in ethanol, cleared in 
beechwood creosote or in eugenol, and mounted in Canada balsam. Thick transverse 
hand-cutting sections of the proglottides were stained following equivalent procedure. 
Two scolices and pieces of strobilas were embedded in paraffin, transversely sectioned 
at 12-15 um, stained with Weigert’s hematoxylin and counterstained with 1% eosin B 
and mounted in Canada balsam. Eggs were mounted in distillated water for drawing. 
Two scolices were prepared for scanning electron microscopy (SEM), they were 
dehydrated through a gradual series of ethanol, then put in amyl acetate, critical point 
dried and sputtered with gold and photographed with a Zeiss DSM 940 A SEM. 

Rudolphiella szidati sp. n. type material was deposited at the Helminthological 
Collection of the “Museo Argentino de Ciencias Naturales Bernardino Rivadavıa”, 
Buenos Aires, Argentina (MACN), and at the Natural History Museum, Geneva, 
Switzerland (MHNG). We also studied syntypes of R. piranabu (Woodland, 1934) 
BMNH 1964.12.15.101.107 and syntypes of R. myoides (Woodland, 1934) BMNH 
1964.12.15.108.110, (Amaz 74.3), both parasites from Pinirampus pirinampu (Spix, 
1829); syntypes of À. piracatinga (Woodland, 1935) BMNH 1964.12.15.206-208, 
(Amaz 40) from Calophysus macropterus (Lichtenstein, 1819) from the British 
Museum of Natural History, London (BMNH). Amaz = field numbers of Woodland’s 
material. 


RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 83 


All measurements are given in micrometres, unless otherwise stated, with the 
range followed by the mean (m) and the number of measurements (n) in parentheses. 
MT = type material. Illustrations were made with the aid of a camera lucida. 


RESULTS 


Rudolphiella Fuhrmann, 1916 
Syn. Amphilaphorchis Woodland, 1934 


Diagnosis: Proteocephalidea, Monticelliidae, Rudolphiellinae. Worms of small 
or medium size; wrinkle collar-like metascolex; suckers uniloculate with developed 
internal circular musculature in their distal margin; apical glandular cells arranged in a 
cross situated between the suckers and the apical tegument; internal longitudinal 
musculature developed; testes cortical in one layer; vagina posterior or anterior to cirrus 
pouch, when anterior ventrally overlapping the cirrus pouch; genital pores irregularly 
alternating; ovary medullar with outgrowths in cortex, strongly lobulate; vitelline 
follicles cortical, ventral, with a tendency of posterior concentration; uterine primor- 
dium and lateral branches medullar; eggs with elongated shell and with embryophores 
bearing two elongated polar projections; all species parasites of catfishes of the family 
Pimelodidae. Type species: Rudolphiella lobosa (Riggenbach, 1895) 


Rudolphiella szidati sp. n. Figs 1, 3-5, 8, 11 


Host: Luciopimelodus pati (Valenciennes, 1840) (= Pimelodus pati), common name: patt. 
Material studied: Argentina, Provincia de Corrientes, Ciudad de Corrientes, Puerto Italia (Parana 
river), holotype MHNG: 26251 INVE, 30.07.1997, paratypes MHNG: 26252 - 26256 INVE, 
30.07. to 01.08.1997; Argentina, Provincia de Chaco, Pte General Belgrano (Parana river) (27° 
27'S, 58° 50°W), 8 paratypes MACN: 392/1-8; Argentina, Provincia de Buenos Aires, Puerto de 
Buenos Aires (de la Plata river) (34° 37°S, 58° 22°W), MACN 392/9-10, 2 paratypes, 06.06.1995; 
other material: Argentina, Provincia de Corrientes, Ciudad de Corrientes, Puerto Italia (Parana 
river), 24667 - 24674, 27235 INVE, 30-31.07.1997 and MACN 392/11-15. 

Site of infection: anterior portion of intestine and rarely in first portion of 
medium intestine. 

Prevalence: 100%, 62 fishes examined. 

Intensity: 2-52; m = 20 

Etymology: the new species honours Prof. Lothar Szidat (Buenos Aires). 
DESCRIPTION (based on 30 specimens and measurements on 8 specimens): 

Monticelliidae, Rudolphielliinae. Testes cortical, situated dorsally and surroun- 
ding laterally and ventrally the longitudinal muscular bundles, reaching but not over- 
lapping the vitelline follicles (Fig. 4); ovary medullar with projections into the cortical 
parenchyma (Fig. 5); uterus medullar (uterine stem and lateral branches). Vitelline 
follicles cortical in 2 ventral bands (Figs 1, 4-5). 

Medium size worms, flattened dorso-ventrally, total length 16-27 mm (m = 22, n 
= 8). Strobila with wrinkles and furrows, acraspedote, comprising about 22-37 (m = 28, 
n = 8) proglottides, fast maturation (10-18 immature proglottides, 4-6 mature pro- 
glottides and 6-10 gravid proglottides). 


84 


ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER 


ROS A) En ae = 2 g & 


A 


ci 28 68 < EN 7 ; > rte i 2 0. a 
= SERI GE Sa DIL > SER = Na = oS 
SR OD PRES o Ÿ DU = = 
Te POS — 


D> 


a 


RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 85 


Wrinkle collar-like metascolex (in the sense of de Chambrier & Paulino, 1997) 
(Fig. 11), encircling the anterior part of the scolex, 1735-2970 in diameter. Metascolex 
with apical elongated bottle-shape glandular cells with granular inclusions, arranged in 
a cross situated between the suckers and the apical tegument (Fig. 3a); 4 slightly oval 
uniloculate suckers. 395-595 (m = 490, n = 19) long and 345-560 (m = 415, n = 19) 
wide; presence of internal circular musculature on distal margin of suckers (Fig. 3b): 
suckers usually hidden by the collar-like metascolex. The suckers are visible when the 
scolex is fixed outside the gut. 

Proliferation zone 990-2970 (m = 2005, n = 8) long, 625-1050 (m = 845, n = 8) 
wide. Immature proglottides 130-530 (m = 310, n = 26) long, 695-1485 (m = 1080, n = 
26) wide; mature proglottides 380-1110 (m = 610, n = 34) long, 925-1780 (m = 1330, n 
= 34) wide. Gravid proglottides 495-1500 (m = 965, n = 37) long, 775-1895 (m = 1425, 
n = 37) wide. Last proglottis 2145 long, 990 wide. 

Internal longitudinal musculature developed, forming a thick bundle of thin 
muscular fibres (Figs 4-5). Osmoregulatory canals thick-walled, with anastomoses in 
the last third of proglottis. Ventral canals interconnected posteriorly to ovary (Fig. 1). 
Ventral canals 20-45 in diameter and dorsal canals 8-30 in diameter. A thin lateral 
ventral secondary duct of ventral osmoregulatory canal opens on each posterior side of 
the proglottis, forming a vesicle before ending in the tegument. This structure was also 
observed in R. lobosa by Riggenbach (1896). 

Testes spherical, 50-70 in diameter, 289-365 (m = 325, n = 6), one-layered, in a 
compact single field, located in the cortical parenchyma, dorsal, lateral and ventral in 
part, not interrupted at cirrus pouch level; overlapping the anterior margin of the ovary 
and laterally reaching the posterior end of the proglottis (Fig. 1). 

Genital pores alternating irregularly, in 20-46% (m = 30%, n = 34) anterior of 
proglottis length. Ejaculatory duct coiled. Cirrus pouch thin-walled, 230-330 (m = 270, 
n = 34) long, 60-110 (m = 85, n = 34) wide. Cirrus pouch length occupying 15-32% (m 
= 23%, n = 34) of proglottis width. Vas deferens bulky and very coiled, sometimes 
reaching body midline, usually reaching anterior margin of mature proglottis, 
occupying aproximately one third of mature proglottis length (Fig. 1). 

Vagina thick- walled, forming 1-2 loops near the ovary. Vagina posterior (77%) 
or anterior (23%) to cirrus pouch; when the vaginal duct is anterior, ventrally 
overlapping cirrus pouch, not surrounding the coiled vas deferens as is common. Ovary 
medullar, bilobed, strongly lobulate with dorsal and ventral outgrowths, only dorsal 
projections reach the cortex. Ovary occupying 43-68% (m = 53%, n = 34) of proglottis 
width (Fig. 1). 


Fics 1-2 
I. Rudolphiella szidati sp. n., holotype 26251 INVE, pregravide proglottis, dorsal view. 
2. Rudolphiella lobosa (Riggenbach, 1896), syntype-material, 43/43, pregravide proglottis, 
ventral view (coll. Zoological Institute, Neuchatel). The vitelline follicles on the right side are not 
entirely represented. Scale-bar, 500 um. 


ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER 


86 


iew of scolex, detail of the apical 


INVE, apical v 


= 
D 


2467 


3a. 


idati Sp. n. 


Be 


b. Rudolphiella s 


portion showing the granular cells in a cross shape situated below the tegument. 3b. 27235 INVE, 


parasagittal 


FIGS 3a- 


circular musculature in their distal margin. 


oO 


section of scolex showing the internal 


Scale-bar, 500 um. 


RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 87 


ao ae 


os 


Fics 4-5 


Rudolphiella szidati sp. n., paratype, 26253 INVE, transverses sections of a pregravide proglottis. 
4. Sections at ovary level; 5. Sections at posterior part of proglottis. Scale-bar, 1000 um. 


Vitelline follicles cortical in 2 ventral bands, sometimes reaching anterior and 
posterior margin of proglottis, with a tendency to a posterior concentration of follicles, 
uninterrupted by cirrus pouch and vagina (Figs 1, 4-5). 

Uterine primordium medullar originating from a cylindrical mass of chromophil 
cells. Lumen appearing in first mature proglottides. Medullar uterine branches up to 
70% (m = 34%, n = 37) of gravid proglottis width, with 9-17 lateral branches opposite 
to cirrus pouch and 7-16 on cirrus pouch side. 

Eggs with elongated polar projections aproximately of equal size (Fig. 8); thin, 
hyaline and spindle-drop shaped external shell. Outer envelope 128-167 (m = 140, n = 
12) long, 20-23 (m = 21, n = 12) wide. Inner envelope consisting in bilayered embryo- 
phore, with nucleate envelope 16-21 (m = 20, n = 12) long, 10-16 (m = 14, n= 12) wide 
and external layer 40-50 (m = 45, n = 12) long, 18-21 (m = 20, n = 12). Oncosphere 
slightly oval, 10-13 (m = 11, n = 12) long, 7-13 (m = 10, n = 12) wide; oncosphere 
hooks, 3-6 (m = 4, n = 17); According to Swiderski (1994), we interprete the described 
structures as: 1. Shell; 2. Outer envelope; 3. Inner envelope consisting in bilayered 
embryophore with external layer much bigger than nucleate envelope; 4. Oncospherical 
membrane rarely visible; 5. Oncosphere. 


88 ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER 


Rudolphiella lobosa (Riggenbach, 1895) Fuhrmann, 1916 Fig. 2 
syn. Corallobothrium lobosum Riggenbach, 1895; Ephedrocephalus lobosum (Riggenbach,1895) 
Mola, 1906. 


Host (according to Riggenbach, 1895): Pimelodus pati (= Luciopimelodus pati (Valenciennes, 
1840), common name: pati. 


Material examined: Paraguay, Paraguay river, type-material, slides 43/43-44 (Collection of the 
Institute of Zoology, Neuchatel, deposited in the MHNG), collected by Ternetz, 01-02.1894. 


Site of infection: intestine. 


DESCRIPTION: 


Strobila acraspedote, with wrinkles and furrows; worms of medium size 17-22 
mm long, with fast maturation. Scolex with apical glandular cells arranged in a cross, 
situated between the suckers and the apical tegument, presence of a well developed 
metascolex, 1025-1350 wide. Suckers uniloculate 205-245 in diameter, with free distal 
muscular sphincter. Proliferation zone about 800 long. Longitudinal muscular fibers 
developed all around the proglottis. 

Testes in one layer in transverse sections, in a single field, sphaerical 50-65 in 
diameter, 194-219 (m = 206; n = 3) (53 in original figure of Riggenbach (1896), 150- 
200 according to Riggenbach (1896), 220 according to Fuhrmann (1916)) in number. 
Testes in cortical parenchyma, dorsal, lateral and ventral in part, not interrupted at 
cirrus pouch level; overlapping anterior margin of ovary and laterally reaching posterior 
end of the proglottis (Fig. 2). 

Genital pores alternating irregularly, in 21-32% (n = 6) of proglottis length. 
Cirrus pouch thin-walled, 170-205 long. Cirrus pouch length occupying 16-22% (n= 4) 
of proglottis width. Vas deferens not reaching body midline in mature proglottis. 

Vagina posterior (53%) or anterior (47%) to cirrus pouch, with inconspicuous 
terminal muscular sphincter. Ovary medullar, bilobed, strongly lobed and bearing 
dorsal projections penetrating the cortex; occupying 60-72% of proglottis width. 
Vitelline follicles distributed cortically and ventrally, in two lateral bands, sometimes 
reaching anterior and posterior margin of proglottis, with a tendency of posterior 
concentration, uninterrupted by cirrus pouch and vagina. 

Uterine primordium like a medullar cylindrical mass of chromophil cells. 
Lumen appearing from the first mature proglottides until formation of eggs. Uterine 
branches occupying up to 56% of proglottis width, with 8-12 lateral branches on each 
side, lor 2 layered. Eggs with two polar projections. 

Remarks: The new species belongs to Rudolphiella Fuhrmann, 1916 based on 
the cortical distribution of the testes and vitelline follicles, on the medullar position of 
uterus, as well as on the medullar and partially cortical location of the ovary (Rego, 
1994). Four species of Rudolphiella are present: R. lobosa (Riggenbach, 1895) recorded 
in Luciopimelodus pati: R. myoides (Woodland, 1934) and R. piranabu (Woodland, 
1934), both parasites of Pinirampus pirinampu; Rudolphiella piracatinga (Woodland, 
1935), parasite of Calophysus macropterus. Rudolphiella cf. lobosa de Chambrier & 
Vaucher, 1999 parasite of Megalonema platanum is also considered. 


RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 89 


Rudolphiella szidati sp. n. differs from all described species in the genus by the 
size of metascolex and by the testes number (Table 1). Furthermore, in R. szidati, (with 
R. lobosa and R. cf. lobosa) testes laterally reaching the posterior end of proglottis, 
while in R. myoides and R. piranabu the number of testes decreases and they do not 


Fics 6-10 


Rudolphiella spp., Eggs drawn in distilled water. 6. Rudolphiella piranabu (Woodland, 1934), 
25129 INVE; 7. Rudolphiella myoides (Woodland, 1934), 24712 INVE; 8. Rudolphiella szidati 
sp. n., 24670 INVE; 9. Rudolphiella piracatinga (Woodland, 1935), 19650 INVE; 10. 
Rudolphiella cf. lobosa, 22352 INVE (de Chambrier & Vaucher, 1999, fig. 95). Several eggs are 
not totally ripe. Abbreviations: eb = bilayered embryophore, oe = outer envelope, on = 
oncosphere, s = shell. Scale-bar, 50 um 


90 ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER 


reach the posterior end of proglottis (de Chambrier & Vaucher, 1999; Pavanelli & 
Machado dos Santos, 1992; Riggenbach, 1895; Woodland, 1934, 1935). 

Within the genus Rudolphiella, the eggs have a similar pattern with elongated 
poles, R. szidati (Fig. 8), R. piracatinga (Fig. 9), R. cf. lobosa (Fig. 10) and R. piranabu 
(Fig. 6) possess poles of similar size, but the shape is different in the 4 taxa; in R. 
myoides, one pole is shorter than the other (Fig. 7). The oncospheres of the 4 known 
species are similar in size (9-13), the outer envelope is difficult to compare among 
species based on uncertainties in the descriptions by different authors. 


DISCUSSION 


As we observed some more characters which are shared by all Rudolphiella 
species (i.e. presence of glandular cells within the apex, vagina anterior/posterior, shape 
of eggs), we gave above a new diagnosis of the genus. 

Luciopimelodus pati is a common dweller fish of the Paraguay river, Bermejo 
river, Parana river, Uruguay river, Carcarana river and de la Plata river. The distribution 
of 3 Pimelodidae sharing the same common name (« pati »), Megalonema platanum 
(Günther, 1880), Pinirampus pirinampu (Spix, 1829) and Luciopimelodus pati 
(Valenciennes, 1840) is overlapping (Ringuelet er al., 1967). Since we were unable to 
find the species of parasites described by Riggenbach (1895) (neither R. lobosa nor 
Proteocephalus fossatus), from L. pati, we suspect that the host studied by Riggenbach 
was not Luciopimelodus pati, but possibly Megalonema platanum; this host was studied 
for cestodes by de Chambrier & Vaucher (1999) and it was parasited by Rudolphiella 
cf. lobosa. 

Rego (1975) described Monticellia rugata from Calophysus macropterus in 
Amazonia. This cestode is a mixture of the two species Nomimoscolex piracatinga 
Woodland, 1935 and Monticellia piracatinga Woodland, 1935. The figures 44, 45, 47, 
49 and 50 belong to Monticellia piracatinga and figures 46 and 48 belong to Nomi- 
moscolex piracatinga. This latter species was transfered to the genus Monticellia under 
the name Monticellia amazonica by de Chambrier & Vaucher (1997). «Piracatinga» is 
the vernacular name given to Calophysus macropterus in Amazonia and to 
Luciopimelodus pati in high Parana river in Brazil. Fowler (1951) and Ringuelet er al. 
(1967) placed L. pati into the Parana bassin, but not in the Amazon bassin. We think 
that Woodland could be mistaken in attributing the name piracatinga to L. pati in 
Amazonia. Our recently collected material in Amazonia from Calophysus macropterus 
fits with the type material of the two species described by Woodland (M. piracatinga 
and N. piracatinga) and seems to confirm our hypothesis. Monticellia rugata Rego, 
1975 is thus a junior synonym of Monticellia piracatinga. However, the study of type 
material of M. piracatinga Woodland, 1935 revealed that this species belongs to the 
genus Rudolphiella, as it is confirmed by the recent collected material from Calophysus 
macropterus in the Amazon and becomes Rudolphiella piracatinga (Woodland, 1935) 
comb. nov. 

So far, numerous proteocephalideans possess unicelullar glands in the scolex (de 
Chambrier ef al., 1992, 1996; de Chambrier & Vaucher, 1997, 1999; Gil de Pertierra, 


RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 9] 


Fic. 11 
Scolex, Rudolphiella szidati sp. n., 24669 INVE. Scale-bar, 200 um. 


1995; Gil de Pertierra & Viozzi, 1999; Scholz et al., 1998, 1999; Stoitsova et al., 1995). 
These glands are also present in all known species of Rudolphiella in high number and 
arranged in a cross, with granular inclusions. Zd“arska & Nebesarova (1999) give a 
detailed description of the unicellular glands situated under the apex of the scolex in 
Proteocephalus macropterus. 

The presence of R. myoides and R. piranabu was confirmed in the host Pini- 
rampus pirinampu from Amazon river by one of the authors (A. de Chambrier), and R. 
piranabu was confirmed by Pavanelli & Machado Dos Santos (1992) in the same host 
from Parana river, Brazil. 

Brooks (1984, 1995) already demonstrated, in phylogenetic analyses based on 
morphological characters, the monophyly of three Rudolphiella species (R. lobosa, R. 
myoides and R piranabu) based on the presence of elongated egg. However, his 1995 
analysis showed some homoplasious characters and is consequently somewhat less 


ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER 


02 


yonod SNIIIO JIM uosLedwos ur PUISEA JO UONISOd 4 


(T6GI) SOIUES SOP Opeyse N © [[[oUBAR 0) SUIPIO99P xx 
(SEGI) PUPIPOOMA 01 SUIPI099P Fununow UL P9PAOJSIP x 


J0119Jsod 9,87 J0119JSOA 9,€€ J0119)SOd % [8 JOLI9]SOd 09€ IOL9]SOd %LL «xx UONISOA BUISEA 

wu ()] ULI || wu () | wu 9 ww ZT-LI 77 = u ‘uw /7-9] SU] [EJOL 

LOSC SI SI SI SE LE=CC smoysold Jo ‘qu 

OI-+ zı-8 L1-6SPPHAPAIP QUI ‘qu 

€81-081 001 08) 001 + 0502 900 EL GITE Hol 2 STEWIE EE GT JOQUUNU 59)59) 

Iz = u 97-81 LOUE EAN Og = uU 97-07 aod russ % 

85 = u 79-45 C9 = TL-09 go = u 89-€Y AIVAO % 

COS UE SIC CO) COUT ESC Jd % 

SPE-COE SVC-SOC SOS-CGE SIOHONS 9 

OOLI=ST6 06S 089-085 x OYE-OFE OSEI-STOI OL6T-SELI X9[OOSEJOLI 9 

vurled UOZLUIV UOZLWV UOZEUUVY vuried Buried USE 
wnupjpjd ndumuntd ndwpund sn]pydoso4apuı 

DULIUO]D3IAW sndwp.nug sndwpsuid snskydo]p) „upd snpojoung,, upd snpojaundoiny SISOH 

(6661 ‘At'Y¥O 2P) sod us sodAjuAs sod {jus sadAJuAs 
DS0Q0] PM sapioXW ‘Y nqpupald ‘y psuyvoviid "y DSOgO] ‘y ‘u ‘ds yppı2s ‘y  D]]21djopny snusy 


‘satoads n7]21Ydjopny zo Sainseaut JAanemedwo,) ‘| AIAVL 


RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 93 


supported. We observed that all Rudolphiella species shown some more synapo- 
morphies: presence of glandular cells with granular inclusions arranged in a cross 
situated between the suckers and the tegument, eggs with elongated shell, embryo- 
phores bearing two polar projections, vitellines follicles cortical in two ventral bands 
situated sub-laterally, vagina anterior/posterior. Those synapomorphies shared by all 
Rudolphiella species emphasize the homogeneity of this group and enforced the 
Brooks’ opinion on the monophyly of this genus. 

Stewart (1986) stated that Luciopimelodus pati, Pinirampus pirinampu and 
Calophysus macropterus belong to a monophyletic group called the «Calophysus 
group», within the Pimelodidae, opinion confirmed by Lundberg ef al. (1991) and more 
recently by de Pinna (1998). On the other hand, Megalonema shares some charac- 
teristics with the Calophysus group, and other groups closely related to Diplomystidae 
(primitive catfishes). In the light of our studies, the presence of Rudolphiella ct. lobosa 
(de Chambrier & Vaucher, 1999) from Megalonema platanum confirms the relationship 
of M. platanum with the Calophysus group. 

In conclusion, the sole genus Rudolphiella is widespread among the genera of 
the «Calophysus group» we studied, including Megalonema platanum. It is, worthwhile 
to emphasize that it could be a likely case of coevolution between related species of 
hosts and their parasites. 


ACKNOWLEDGEMENTS 


We are grateful to Dr. C. Vaucher for helpful discussions and for inviting the 
first author to the Natural History Museum, Geneva; to the rewievers for constructive 
critisism; to the Municipal Departement of Cultural Affaires, Geneva, Switzerland, for 
supporting her stay at the Department of Invertebrates of the Museum in 1998. Her visit 
was also supported by the Secretaria de Recursos Naturales y Desarrollo Sustentable de 
la Nacion, Buenos Aires, Argentina. We thank the staff of the Centro de Ecologia 
Aplicada del Litoral, Corrientes, Argentina for assisting our stay in the field during 
1997 and particularly to Mr. L. Benetti, Prof. J. J. Neiff, Dr. M. Hamann and Dr. A. 
Kehr. We are thankful to Dr. E. A. Harris, at the British Museum of Natural History, for 
the loan of specimens. 


REFERENCES 


Brooks, D. R. 1995. Phylogenetic hypothesis, cladistic diagnoses, and classification of the 
Monticelliidae (Eucestoda: proteocephaliformes). Revista Brasileira de Biologia 55: 
359-367. 

Brooks, D. R. & RASMUSSEN, G. 1984. Proteocephalid cestodes from Venezuelan catfish, with a 
new classification of the Monticelliidae. Proceedings of the Biological Society of 
Washington 97: 748-760. 

DE CHAMBRIER, A. & PAULINO R. C. 1997. Proteocephalus joanae n. sp. (Eucestoda: Proteo- 
cephalidea), a parasite of Xenodon neuweidi (Serpentes: Colubridae) from South 
America. Folia Parasitologica 44: 289-296. 

DE CHAMBRIER, A., SCHOLZ, T. & VAUCHER, C. 1996. Tapeworms (Cestoda: Proteocephalidea) of 
Hoplias malabaricus (Pisces: Characiformes, Erythrinidae) in Paraguay: description of 
Proteocephalus regoi n. sp., and redescription of Nomimoscolex matogrossensis. Folia 
Parasitologica 43: 133-140. 


94 ALICIA A. GIL DE PERTIERRA & ALAIN DE CHAMBRIER 


DE CHAMBRIER, A. & VAUCHER, C. 1997. Revision des cestodes (Monticelliidae) décrits par 
Woodland (1934) chez Brachyplatystoma filamentosum avec redéfinition des genres 
Endorchis Woodland, 1934 et Nomimoscolex Woodland, 1934. Systematic Parasitology 
811322119233: 

DE CHAMBRIER, A. & VAUCHER, C. 1999. Proteocephalidae and Monticelliidae (Eucestoda: 
Proteocephalidea) parasites de poissons d’eau douce au Paraguay, avec descriptions d’un 
genre nouveau et de dix especes nouvelles. Revue suisse de Zoologie 106: 165-240. 


DE CHAMBRIER, A., VAUCHER, C. & RENAUD, F. 1992. Etude des caracteres morpho-anatomiques 
et des flux géniques chez quatre Proteocephalus (Cestoda: Proteocephalidae) parasites de 
Bothrops jararaca du Bresil et description de trois especes nouvelles. Systematic Para- 
sitology 23: 141-156. 

DE PINNA, M. C. C. 1998. Phylogenetic relationships of Neotropical Siluriformes (Teleostei: 
Ostariophysi): Historical overviews and synthesis of hypotheses (pp. 279-330). In: 
Malabarba, L. R., Reis, R.E., Vari, R.P., Lucena, Z. M. S. & Lucena, C. A. S. (eds). 
Phylogeny and classification of Neotropical fishes. Part 3. Siluriformes. Edipucrs, Porto 
Alegre, Brasil: 279-330. 

Fow er, H. W. 1951. Os peixes de agua doce do Brasil, vol. I. Archivos de Zoologia do Estado 
do sao Paulo 6 : 405-625. 


FUHRMANN, O. 1916. Eigentiimliche Fischcestoden. Zoologischen Anzeiger 46: 385-398. 


GIL DE PERTIERRA, A. A. 1995. Nomimoscolex microacetabula n. sp. y N. pimelodi n. sp. (Ces- 
toda: Proteocephalidae) paräsıtos de Siluriformes del Rio de la Plata. Neorröpica, 41: 
19-25. 


GIL DE PERTIERRA, A. A. & VIOZZI, G. P. 1999. Redescription of Cangatiella macdonaghi (Szidat 
& Nani, 1951) comb. nov. (Cestoda: Proteocephalidae) a parasite of the Atheriniform fish 
Odontesthes hatcheri (Eigenmann, 1909) from the Patagonian Region of Argentina. 
Neotropica 45: 13-20. 

LANGERON, M. 1949. Précis de Microscopie, 7° éd., Masson & Cie, Paris, 1429 pp. 

LUNDBERG, J. G., MAGO-LECCIA, F. & Nass, P. 1991. Exallodontus aguanai, a new genus and 
species of Pimelodidae (Pisces: Siluriformes) from deep river channels of South America 
and delimitation of the subfamily Pimelodinae. Proceedings of the biological Society of 
Washington 104(4): 840-869. 

Reco, A. A. 1975. Estudos de cestöides de peixes do Brasil. 2° nota: revisäo do género 
Monticellia La Rue, 1911 (Cestoda, Proteocephalidae). Revista Brasileira de Biologia 35: 
567-586. 

Reco, A. A. 1994. Order Proteocephalidea Mola, 1928 (pp. 257-293). In: Khalil, L. F., Jones, A. 
& Bray, R. A. (eds). Keys to the Cestode Parasites of Vertebrates. Wallingford: CAB 
International. 

RIGGENBACH, E. 1895. Beiträge zur Kenntnis der Taenien der Süsswasserfische. Centralblatt für 
Bakteriologie und Parasitenkunde 18 (20/21): 609-613. 

RIGGENBACH, E. 1896. Das genus Ichthyotaenia. Revue suisse de Zoologie 4: 165-276. 


RINGUELET, R. A., ARAMBURU, R. H. & ARAMBURU, A. A. 1967. Los peces argentinos de agua 
dulce. Comision de Investigacion Cientifica. Provincia de Buenos Aires, La Plata, 
602 pp. 

PAVANELLI, G. C. & MACHADO Dos SANTOS, M. H. 1992. Goezeella agostinhoi n. sp., e Monti- 
cellia loyolai n. sp., cestöides proteocefalideos parasitas de peixes pimelodideos do Rio 
Parana, Parana, Brasil. Revista Brasileira de Parasitologia Veterinaria 1(1): 45-50. 

SCHOLZ, T., DRABEK, R. & HANZELOVA, V. 1998. Scolex morphology of Proteocephalus tape- 
worms (Cestoda: Proteocephalidae), parasites of freshwater fishes in the Palaeartic 
Region. Folia Parasitologica 45: 27-43. 

SCHOLZ, T., ZD'ARSKA, Z., DE CHAMBRIER, A. & DRABEK, R. 1999. Scolex morphology of the 
cestode Silurotaenia siluri (Batsch, 1786) (Proteocephalidae: Gangesiidae), a parasite of 
European wels (Silurus glanis). Parasitology Research 85: 1-6. 


RUDOLPHIELLA SZIDATI SP. N., PARASITE OF LUCIOPIMELODUS PATI FROM ARGENTINA 95 


STEWART, D. J. 1986. Revision of Pimelodina and description of a new genus and species from 
the Peruvian Amazon (Pisces: Pimelodidae). Copeia 3: 653-672. 

STOITSOVA, S., GEORGIEV, B., DRACHEVA, R. & VINAROVA, M. 1995. Ultrastructural and cyto- 
chemical demonstration of two types of scolex glands in Proteocephalus osculatus 
(Cestoda, Proteocephalidea). Comptes rendus de l'Académie bulgare des Sciences 48(8): 
97-99. 


SWIDERSKI, Z. 1994. Ultrastructure of Infective Eggs of Proteocephalus longicollis (Zeder, 1800) 
(Cestoda, Proteocephalidea). Proceedings of the 13th International Congress on Electron 
Microscopy, Paris, 3b: 1427-1428. 

WOODLAND, W. N. F. 1934. On the Amphilaphorchidinae, a new subfamily of proteocephalid 
cestodes, and Myzophorus admonticellia, gen. et n. sp., parasitic in Pirinampus spp. from 
the Amazon. Parasitology 26: 141-149. 

WOODLAND, W. N. F. 1935. Some more remarkable cestodes from Amazon siluroid fish. Para- 
sitology 27: 207-225. 

ZbD ARSKA, Z & NEBESAROVA, J. 1999. Distribution and ultrastructure of two types of scolex 


gland cells in adult Proteocephalus macrocephalus (Cestoda, Proteocephalidea). Parasite 
6: 49-56. 


È 


| | | 
RA ABR Talarico 


REVUE SUISSE DE ZOOLOGIE, 107 (1): 97-106; mars 2000 


Two new species of the genus Suffasia from Sri Lanka 
(Araneae: Zodariidae) 


Suresh P. BENJAMIN! & Rudy JOCQUE? 

l'Institut für Natur-, Landschafts- und Umweltschutz der Universität Basel (NLU), 
Abteilung Biologie, St. Johanns-Vorstadt 10 
CH-4056 Basel, Switzerland. 

2 Koninklijk Museum voor Midden-Afrika, B-3080 Tervuren, Belgium. 


Two new species of the genus Suffasia from Sri Lanka (Araneae: 
Zodariidae). - Two new species of the genus Suffasia Jocqué, 1991 are 
described. S. mahasumana sp. n. 1s known from both sexes and is related to 
S. tumegaster Jocqué, 1992. S. attidiva sp. n., known only from two female 
specimens, may be related to S. tigrina (Simon, 1893). S. attidiya Sp. n. is 
found in diverse habitats, S. mahasumana sp. n. is confined to cloud forests 
in the central highlands of Sri Lanka. This is the first record of Suffasia 
from Sri Lanka, other species in this genus are known only from southern 
India and Nepal. 


Key-words: Araneae - Zodariidae - Suffasia - tropical montane cloud forests 
- Sri Lanka. 


INTRODUCTION 


The genus Suffasia was established for two species from southern India. The 
type species, S. tigrina (Simon, 1893) from Kodaikanal, Tamil Nadu, and an undescri- 
bed species from the same locality (Jocqué, 1991). S. tumegaster Jocqué, 1992, from 
Kathmandu, Nepal, was added later. Considering the affinities of the faunas of India 
and Sri Lanka it was to be expected that members of Suffasia or at least of closely 
related genera might be discovered on the island. 

During recent field work in Sri Lanka two undescribed species were collected. 
The first one is from the Knuckles Range, consisting of a number of remnant patches of 
primary tropical montane cloud forests, a type of vegetation that was once common in 
the central highlands of Sri Lanka. Most of these forests were cleared for tea plantations 
during the British colonial period. The second species was collected in fragmented 
marshland situated on the outskirts of Colombo. On both localities the specimens were 
obtained by beating shrubs and small trees which is an unusual way to collect zodariids 
as they are considered soil dwelling spiders except for the representatives of the 
Storenomorphinae (Jocqué, 1991). In the Bellanwila-Attidiya sanctuary a marshy area 
near Colombo, shrubs are scattered around shallow water ponds, marshes and sea- 


Manuscript accepted 21.10.1999 


98 SURESH P. BENJAMIN & RUDY JOCQUE 


sonally flooded grassland. A female of the second new species was collected by the 
same method in Kalugala, Labugama Forest Reserve, a fragmented remnant of tropical 
lowland rain forest, some 40 km away from the former locality. 


METHODS 


Structures were examined in temporary mounts embedded in glycerine. Vulvae 
were Cleared with trypsin (0.1% trypsin, 0.1% CaCl,, in 0.05M tris-buffer, pH 7.6). All 
drawings were made with a Nikon Labophot-2 and a Nikon SMZ-U microscopes with 
drawing tube. Measurements are in mm. Structures examined with the scanning elec- 
tron microscope (PHILIPS XL30 FEG ESEM) were critical point dried, stud-mounted 
and sputter coated for observation and photography. Specimens examined are deposited 
in the “Muséum d’histoire naturelle, Geneve” (MHNG) and the “Naturhistorisches 
Museum, Basel” (NMB). 

Abbreviations used in the text and figures: AER anterior eye row; ALE anterior 
lateral eyes; AME anterior median eyes; CD copulatory duct; CF cymbial flange; CO 
copulatory opening; E embolus; FD fertilisation duct; PER posterior eye row; PLE 
posterior lateral eyes; PME posterior median eyes; TA tegular aphophysis. 


Suffasia mahasumana sp. n. Figs 1-7, 13-21 
Holotype 3: Sri Lanka, Central Province, Knuckles Range, Deenston (approximately 7° 19° N, 
80° 51° E), 1100 m, 11 March 1998. Leg. Suresh P. Benjamin (MHNG). 

Paratypes: 1%, 11 March 1998 (MHNG): 14,19 12 March 1998 (NMB); further as holotype. 


Etymology: Named after god Maha-Sumana, protector of the hill country in Sri Lanka. 
Noun in apposition. 


Diagnosis: S. mahasumana is closely related to S. tumegaster; the male can be reco- 
gnised by the absence of a dorsal spike on the palpal tibia, the shape of the dorsal 
cymbial flange which is flat and evenly rounded in the latter, swollen, curved upwards 
and concave in the former; the female differs from that of S. tumegaster by the presence 
of a roughly rectangular plate in the anterior part of the epigyne. 


Description: Male (holotype). Colouration and markings: carapace dark yellow-brown, 
with dark reticulations on anterior part and with U-shaped dark marking in front of 
fovea (Fig. 14). Chelicerae and sternum dark yellow, lighter than carapace. Dorsum of 
opisthosoma uniform darkgrey, venter white, without markings. Legs light yellow with 
dark dorsal markings. AER almost straight, PER slightly procurved, all eyes circular, 
AME 1.5 times their diameter apart from each other and at about the same distance 
from ALE. PME 2 times their diameter apart and about the same distance from PLE. 
ALE = PLE = PME> AME. Clypeus height 6 times the diameter of ALE. Chilum 
present. Chelicerae not fused, with double-tooth on promargin. Labium triangular. 
Sternum sub triangular, with spike-like extensions projecting towards base of coxae. 
Leg formula 4132, 2 spines dorsally on femora I-IV. Tibiae with flattened incised hairs 
(Fig. 16, FIS); and ventral tuft of metatarsal preening brush with chisel-shaped hairs 
(Figs 16, 17, CH; see also Jocqué, 1991: figs 6, 8, 11, 12). Femoral organ (Fig. 20) 
present on each leg. Trichobothrium base with concentric ridges (Fig. 15). 


TWO NEW SUFFASIA FROM SRI LANKA 99 


Palp (Figs 1-4): Tibia with stout, sharp, strongly tapered retrolateral apophysis, 
pointing outwards. Cymbium strongly narrowed in dorsal view, with swollen, upwards 
curved, dorsolateral flange (Fig. 1, CF), extending lateral cymbial concavity, carrying 
some sensorial hairs in superior part; embolus fairly short, stout, originating on 


Measurements: total length: 2.3; carapace length: 1.3; carapace width: 1.0. Legs: 


I Il III IV 
femur 13] 0.9 1.0 iol 
patella 0.2 0.2 0.3 0.3 
tibia 1.0 0.7 0.7 1.0 
metatarsus 1] 0.9 1.0 LS 
tarsus 0.5 0.4 0.4 0.5 
total 3.9 3.1 3.4 4.4 


posterior part of tegulum separated from main part by shallow groove; tegular 
apophysis, short, stout, sharp, pointing out and forwards (Figs 1, 2, TA). 

Female. Colouration and markings: Similar to male but lighter. Different by 
possessing dorsal opisthosomal markings as in Fig. 13; venter white. Palp with conical 
tarsus, longer than tibia (Fig. 19). Morphology further as in male. 

Epigynum and vulva (Figs 5-7): Simple brown plate in anterior part; internal structure 
visible through thin tegument; copulatory openings in front hidden by plate; short 
copulatory ducts lead to thick-walled spermathecae, triangular in dorsal view. 


Measurements: Total length: 3.0; carapace length: 1.3; carapace width: 1.0. Legs: 


I II III IV 
femur 0.8 0.8 0.9 PSI 
patella 0.3 0.3 0.4 0.4 
tibia 0.8 0.7 0.6 0.8 
metatarsus 0.9 0.7 0.8 1.2 
tarsus 0.5 0.4 0.4 0.5 
total 33 2.9 Bell 4.0 


Affinities: Suffasia is defined by the presence of a chilum and promarginal cheliceral 
teeth, dark reticulation (“network pattern” sensu Jocqué, 1992), female palp with a long 
conical tarsus, femoral organ with simple setae on all legs, legs with flattened incised 
hairs and metatarsal preening brush consisting of chisel-shaped hairs (Jocqué, 1991, 
1992). The present species clearly agrees with these characters and can thus be 
attributed to Suffasia. Yet the shape of the male palpal cymbium casts some doubt on 
this attribution. Although one of the main characteristics of Asceua Thorell, 1887 is 
exactly the strongly narrowed cymbium, there are a number of characters that exclude 
the incorporation of the present species in it: representatives of that genus do indeed 


SURESH P. BENJAMIN & RUDY JOCQUE 


100 


Fics 1-4. Suffasia mahasumana sp. n. 1. Male palp, retrolateral view. 2. Ditto, ventral view. 3. 
Ditto, dorsal view. 4. Ditto, prolateral view. CF cymbial flange; E embolus; TA tegular 


apophysis. Scale line: 0.2 mm. 


TWO NEW SUFFASIA FROM SRI LANKA 101 


7 


Fics 5-7. Suffasia mahasumana sp. n. 5. Female epigynum, ventral view. 6. Vulva, ventral view. 
7. Ditto, dorsal view. CD copulatory duct; FD fertilisation duct. Scale line: 0.1 mm. 


102 SURESH P. BENJAMIN & RUDY JOCQUE 


lack teeth, have a uniform dark carapace and lack femoral organs. Yet, both S. maha- 
sumana and S. tumegaster possess an epigynum that is quite different from what has 
been described for Suffasia. It might prove necessary to erect a new genus for these 
species if the male of type species of Suffasia appears equally different which was 
already recognised by Jocqué (1992). 


Distribution: Known only from the type locality. 


Suffasia attidiya sp. n. Figs 8-12 
Holotype 2: Sri Lanka, Western Province, Colombo, Bellanwila-Attidiya (approximately 
6°50°N, 79°54’E), mean elevation 0.6 m asl, 22 February 1998, (MHNG). 

Paratype 2: Sri Lanka, Western Province, Kalugala, Labugama Forest Reserve, 3 August 1996, 
ca. 10 m (NMB). All specimens leg. Suresh P. Benjamin. 

Etymology: Named after the type locality. Noun in apposition. 


Diagnosis: The epigyne of S. attidiva differs from that of S. tigrina by the course of the 
copulatory ducts which run directly inwards in the latter, outwards thence inwards in 
the new species. S. mahasumana is clearly different by the presence of a plate in the 
epigyne. 

Description: Female (holotype). Colouration and markings: prosoma dorsally dark 
yellow-brown, with dark markings in front of fovea (Fig. 12). Chelicerae and sternum 
dark yellow lighter than dorsal prosoma. Dorsum of opisthosoma with markings as in 
Fig. 11, venter white. Legs light yellow with dark dorsal markings. AER slightly pro- 
curved, PER procurved, all eyes circular, AME 1.5 times their diameter apart from each 
other, 0.5 times from ALE. PME 2 times their diameter apart and 2.5 times from PLE. 
ALE = PLE = PME > AME. Clypeus height 6 times the diameter of ALE. Chilum 
present; chelicerae not fused, with double tooth on promargin. Labium triangular. Ster- 
num sub triangular, with spike-like extensions projecting towards base of coxae. Palp 
with conical tarsus, longer than tibia. Flattened incised hairs on tibiae; ventral tuft of 
metatarsal preening brush with chisel-shaped hairs. 

Epigynum and vulva (Figs 8-10): Epigynum simple, anterior sclerotized border with 
CO situated laterally. Thick-walled copulatory ducts straight and close to each other in 
posterior part, leading to small globular rectptacula, with thick walls. FD as in Fig. 9. 
Male. Unknown. 

Affinities: As zodariid genera are mainly diagnosed on male palpal morphology 
(Jocqué, 1991, 1992) the placement of this new species of Suffasia might appear ambi- 
guous. However, the similarities of the epigynum of the present species and of the type 
species are so striking that there is little doubt that they are congeneric. In both cases 
the internal structure is simple with lateral CO, strongly sclerotized CD with a partly 
parallel course and roughly oval spermathecae. 


Fics 8-14. Suffasia attidiva sp. n. (8-12). Suffasia mahasumana sp. n. (13, 14). 8. Female 
epigynum, ventral view. 9. Vulva, ventral view. 10. Ditto, dorsal view. 11. Female opisthosoma, 
dorsal view. 12. Female prosoma and right palp, dorsal view. 13. Female, dorsal view. 14. Male, 
dorsal view. CO copulatory opening; FD fertilisation duct. Scale lines: 0.1 mm (8-10), 2.0 mm 
(11-14). 


103 


ASIA FROM SRI LANKA 


£ 


TWO NEW SUFF 


104 SURESH P. BENJAMIN & RUDY JOCQUE 


Fics 15-18. Suffasia mahasumana sp. n., SEM micrographs. 15. Base of trichobothrium. 16. 
Preening brush on distal metatarsus of leg I, lateral view. 17. Ditto, detail. 18. Tip of leg I, lateral 
view. CH chisel-shaped hairs, FIS flattened incised hairs. Scale lines: 0.001 mm (1), 0.01 mm 
(16-18). 


TWO NEW SUFFASIA FROM SRI LANKA 105 


Fics 19-21. Suffasia mahasumana sp. n., SEM micrographs. 19. Chelicerae and palps of female, 


frontal view. 20. Femoral organ, on leg I. 21. Chemosensitive hair on metatarsus. Scale lines: 
0.003 mm (20), 005 mm (21), 0.1 mm (19). 


106 SURESH P. BENJAMIN & RUDY JOCQUE 


Measurements: Total length: 2.6; carapace length: 1.3; carapace width: 1.0. Legs: 


I II III IV 
femur 0.6 0.5 0.6 0.7 
patella 0.1 0.1 0.2 0.2 
tibia 0.5 0.4 0.4 0.6 
metatarsus 0.6 0.5 0.5 0.8 
tarsus 0.4 0.3 0.3 0.4 
total DID, 1.8 2.0 DOT 


Distribution: Known from Bellanwila-Attidiya sanctuary and Kalugala, Labugama 
Forest Reserve. 


DISCUSSION 


The placement of the two new species in the genus Suffasia is in accordance 
with the current definition of the genus. However the relationships proposed here 
should be re-analysed when additional material, most importantly the male of S. tigrina 
is discovered. 

In his revision of the Zodariidae Jocqué (1991) considered the subfamilies 
Zodariinae and Storeninae to be monophyletic. This hypothesis was based on presumed 
autapomorphies such as the presence of a femoral organ and flattened incised hairs for 
the Zodariinae and chisel-shaped hairs on the metatarsal preening brush for Storeninae. 
The discovery of S. tumegaster, which possesses a combination of all these characters, 
led to an amalgamation of these two subfamilies (Jocqué, 1992). S. mahasumana sp. n. 
which also possesses these three characters, further confirms his combination of both 
subfamilies. 

The discovery of the new taxa extends the previously known distribution (Nepal, 
India) of the genus Suffasia southwards to Sri Lanka. 


ACKNOWLEDGEMENTS 


We thank Dr. Peter Schwendinger (MHNG) for critical review of the manuscript 
and for his encouragement during this study. Part of this work was done in the course of 
the masters thesis of the first author at the University of Innsbruck. He is grateful to Dr. 
K. Thaler for providing research facilities there. We also thank Marcel Diiggelin 
(Raster lab of the Unviersity of Basel) for help with SEM work, Mr. D. Benjamin 
(Colombo) for accompanying the first author on collecting trips to the study area and 
Mr. A. H. Sumanasena (Department of Wild Life Conservation, Colombo) for pro- 
viding a research permit. 


REFERENCES 


JOCQUÉ. R. 1991. A generic revision of the spider family Zodariidae (Araneae). Bulletin of the 
American Muesum of Natural History 201: 1-165. 

JOCQUÉ, R. 1992. A new species and the first males of Suffasia with a redelimitation of the 
subfamilies of the Zodariidae (Araneae). Revue suisse de Zoologie 99: 3-9. 


REVUE SUISSE DE ZOOLOGIE 107 (1): 107-122; mars 2000 


Diversite du zoobenthos dans 47 rivieres du canton de Vaud: 
tendance 1989 - 1997 


Claude LANG 
Conservation de la faune, Marquisat 1, CH-1025 St-Sulpice, Suisse. 
claude.lang@sffn.vd.ch 


Diversity of zoobenthos in 47 rivers of western Switzerland: the 1989 - 
1997 trend. - Three surveys of benthic invertebrates were made between 
1989 and 1997 in 165 sites located in 47 rivers of western Switzerland 
(canton of Vaud). Eight descriptors derived from the total number of taxa 
and the number of taxa intolerant of pollution (Plecoptera, Heptageniidae, 
and Trichoptera with a case) were used to describe the patterns of benthic 
diversity. In the three surveys, the diversity increased with the altitude of 
sampling sites. This trend reflected the upward decrease of human impact 
on the rivers and their watersheds. The diversity increased between 1989 
and 1997 because of better meteorological conditions (i.e. more water in 
the rivers) coupled with the increased efficiency of sewage treatment 
plants. But the diversity remained low in several small rivers, probably 
because of chronic pollutions by pesticides. 


Key-words: diversity - indicator species - river - water quality - zoo- 
benthos. 


INTRODUCTION 


Depuis la signature de la convention sur la diversité biologique a Rio de Janeiro 
en 1992, la conservation et la restauration de la diversité des especes dans l’espace et le 
temps constituent un des buts fondamentaux de l'écologie appliquée (Wright er al., 
1998). Dans le même ordre d'idée, la diversité des communautés d’invertebres qui colo- 
nisent le fond des cours d'eau (zoobenthos) est utilisée depuis le début du siècle pour 
caractériser l’état du milieu: elle diminue en fonction de l'intensité des perturbations 
subies, que celles-ci soient d'origine humaine ou naturelle (Hellawell, 1986). 

Cette diminution de la diversité est particulièrement marquée chez les plé- 
copteres, certains éphéméroptères et les trichopteres à fourreau d’où l’utilisation de ces 
groupes comme indicateurs (Fore ef al., 1996). Dans les rivieres du canton de Vaud par 
exemple, la diversité des plécoptères a fortement diminué entre 1945 et 1982 à basse 
altitude alors qu'elle ne changeait guère dans les rivières de montagne, restées proches 
de l'état naturel (Aubert, 1984). Ce gradient de diversité amont-aval reflète l’augmen- 
tation des impacts d'origine humaine à basse altitude (Lang & Reymond, 1993, 1995). 


Manuscrit accepté le 28.07.1999 


108 CLAUDE LANG 


Entre 1982 et 1997, l'épuration des eaux s'est généralisée et les performances des 
stations d’épuration se sont améliorées dans le canton de Vaud (Fiaux er al., 1998). De 
ce fait, les quantités de matiere organique et de nutriments déversées dans les cours 
d'eau ont diminué. La diversité du zoobenthos devrait donc augmenter en réponse a 
cette amélioration de la qualité chimique de l’eau. 

Il ne faut toutefois pas oublier que le bassin versant et la rivière qui le draîne, 
forment un tout: ce qui affecte l'un, modifie l’autre (Hynes, 1975). Du fait de cette 
relation étroite, le caractere de moins en moins naturel des bassins versants, surtout a 
basse altitude, pourrait empécher la restauration de la diversité, méme si la composition 
chimique de l'eau redevient normale, tout au moins en termes d’éléments majeurs 
(matiere organique, phosphore et azote); les micropolluants constituant en effet un 
problème a part (Corvi & Kim-Heang, 1997; Vioget & Strawczynski, 1997). 

Pour interpréter l'évolution a long terme du zoobenthos, il faut également tenir 
compte de la météorologie (Allan, 1995). Par exemple, une sécheresse de longue durée 
va réduire le débit des cours d'eau et augmenter ainsi la concentration des polluants, 
donc leur impact sur le zoobenthos. Faute d'une dilution suffisante, les capacités d'auto- 
épuration naturelles de certains cours d'eau peuvent étre completement dépassées si 
l’épuration des eaux n’a pas été conçue en fonction de conditions extrêmes. Ces 
dernieres pourraient s’ observer plus fréquemment si le climat venait a changer rapide- 
ment sous l'influence de l’effet de serre. Signalons également que des crues très vio- 
lentes, comme une longue sécheresse, peuvent durablement diminuer la diversité du 
zoobenthos en bouleversant le lit des cours d’eau (Allan, 1995). 

La présente étude analyse l’évolution de la diversité du zoobenthos dans les 
rivieres vaudoises sur la base de trois campagnes de prélevements effectuées entre 1989 
et 1997. Son but consiste a mettre en évidence l’effet sur le zoobenthos des mesures 
d'assainissement prises, ceci en fonction de l’altitude et des conditions météorologiques 
observées au cours de cette période. 


STATIONS ET METHODES 


Le zoobenthos des rivieres vaudoises est étudié depuis 1982 (Lang & Reymond, 
1995). Cependant le réseau de surveillance, sous sa forme actuelle, n'a été mis en place 
qu'à partir de 1989. De ce fait l'analyse présentée ici se concentre sur une période ou les 
données sont parfaitement comparables entre elles: 165 stations de prélevements 
étudiées au cours de trois campagnes effectuées entre 1989 et 1997 dans 47 rivières. 

Les 47 rivières étudiées (Fig. 1) sont réparties en trois groupes d'après la région 
où chacune d'elles prend sa source: (1) le Jura et l'ouest du canton de Vaud, (2) le Jorat 
et le centre du canton, (3) les Préalpes et les Alpes. La Broye, I’ Arbogne et la Mionne 
sont rattachées aux rivieres du Jorat en raison de leur ressemblance avec celles-ci. Dans 
le Jura, 20 rivières et 75 stations de prélèvement ont été visitées en 1990, 1993 et 1996 
(Lang, 1997); dans le Jorat, 15 riviéres et 47 stations visitées en 1991, 1994 et 1997 
(Lang, 1998); dans les Alpes, 12 riviéres et 43 stations visitées en 1989, 1992 et 1995 
(Lang, 1996). Seules les rivières et les stations visitées au cours des trois campagnes 
sont incluses dans la présente étude. 


DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 109 


Fic. | 


Localisation et numéros d'identification des 47 rivieres vaudoises visitées entre 1989 et 1997. 


1 Doye 11 Saubrette 21 
2 B. de Nyon 12 B. Morges 22 
3 Asse 13 Morges 23 
4 Promenthouse 14 Venoge 24 
5 Colline 15 Veyron 25 
6 Cordex 16 Nozon 26 
7 Serine 17 Orbe Dil 
8 Dullive 18 Mujon 28 
9 Aubonne 19 Arnon 29 
10 Toleure 20 Baumine 30 


Talent 
Buron 
Mentue 
Sauteru 
Petite Gläne 
Broye 
Arbogne 
Lembe 
Cerjaule 
Mérine 


31 Bressonne 41 Gryonne 


32 Carrouge 42 P. Gryonne 
33 Grenet 43 Avançon 
34 Mionne 44 Hongrin 
35 Forestay 45 Sarine 


36 Veveyse 46 Flendruz 

37 B. Clarens 47 Torneresse 
38 B. Montreux 1 - 20 Jura 
39 Tinière 21 - 35 Jorat 
40 Grande Eau 36 - 47 Alpes 


Les 165 stations de prélèvements étudiées ont été choisies parce qu’elles pré- 
sentent des caractéristiques communes qui les rendent comparables entre elles: pour la 
plupart, elles sont localisées dans la zone à truite de Huet et le rithron d’Illies, donc 


110 CLAUDE LANG 


TABLEAU 1. Calcul de la valeur de l'indice RIVAUD à partir du nombre total de taxons (NT) et du 
nombre de taxons sensibles aux pollutions (NTS). La valeur de RIVAUD se lit à l'intersection de 
la ligne renfermant la valeur de NT et de la colonne ou se trouve la valeur de NTS. Exemple: NT 
= 19, NTS = 7, RIVAUD = 11. Source: Lang & Reymond (1995). 


NTS 
NT 0 I 2 3 4 5 6-7 8-9 10-11 12-13 14-25 
1-4 I 2 3 4 5 - - - - - - 
5-8 2 3 4 5 6 7 8 9 - - - 
9-12 3 4 5 6 7 8 9 10 ll 12 - 
13-16 4 5 6 7 8 9 10 ll 2 13 14 
17-20 5 6 7 8 9 10 1] 12 13 14 15 
21-24 6 7 8 9 10 1] 12 3 14 15 16 
25-28 Ti 8 9 10 1] 12 13 14 [es 16 IM 
29-32 8 9 10 1] 12 13 14 15 16 17 18 
33-36 9 10 Il 12 13 14 15 16 17 18 19 
37-50 10 1] 12 13 14 15 16 17 18 19 20 


Qualité biologique mauvaise (RIVAUD | - 5), faible (6 - 9), médiocre (10 - 11), moyenne (12 - 
14), bonne (15 - 20). 


dans une écorégion relativement homogene (Ribaut, 1966); toutes renferment des 
surfaces de cailloux balayées par le courant (zone de rapides) d’au moins 20 m? ce qui 
permet de prélever toujours dans le méme substrat; aucune n'est directement exposée 
aux rejets polluants d’une station d'épuration ou d'un important égout: elles représentent 
donc la situation générale de la rivière en l’absence de pollutions ponctuelles bien 
marquées. 

Dans le Jura et le Jorat, chaque station est visitée à deux reprises chaque année 
d'étude: la première fois en janvier-février, la seconde fois en mars-avril, c'est-à-dire 
pendant l’étiage d’hiver et avant la crue de printemps. Dans les Alpes, la premiere 
campagne s'effectue en février-mars pendant l'étiage hivernal, la deuxième en juin 
après la crue de printemps et la troisième en septembre en période de basses eaux. Trois 
visites par année sont nécessaires dans les Alpes. Certaines stations ne sont en effet pas 
accessibles en hiver à cause de la neige et la glace, en juin à cause des hautes eaux et en 
septembre parce que le niveau d’eau est trop bas pour prélever. De plus, du fait du 
caractère torrentiel de ces cours d'eau, l’abondance de la faune y est souvent très faible 
ce qui fait qu’une visite peut ne pas être représentative. 

Au cours de chaque visite, six coups de filet sont donnés dans six différentes 
zones de cailloux (rapides) de la station, correspondant chacune à une surface prélevée 
d'environ 0.1 m2. Le filet est posé sur le fond, son ouverture (20 cm x 20 cm) face au 
courant, de manière à ce que celui-ci y entraîne les invertébrés délogés en remuant le 
substrat avec le pied. Tous les invertébrés récoltés dans ces six coups de filet consti- 
tuent un prélèvement qui est immédiatement conservé sur le terrain dans du formol 
DL 

En laboratoire, les invertébrés, séparés des sédiments et débris végétaux par des 
tamisages successifs (Reymond, 1995), sont identifiés et comptés jusqu’au niveau du 
genre, de la famille ou de la classe selon les groupes (Annexe 1). La liste combinée des 


DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 11] 


taxons présents chaque année dans chaque station est ensuite dressée a partir des deux 
ou trois (Alpes) prélevements effectués dans chacune d’elles; cependant, le méme taxon 
observé dans les deux ou trois prélèvements de la même station n’est compté qu’ une 
seule fois. Toutes les analyses présentées dans cet article sont basées sur la liste 
combinée des taxons présents la méme année dans la méme station. Cette approche 
rend les comparaisons entre stations et entre années plus fiables en diminuant l'impact 
d’evenements exceptionnels (breves crues avant le prélevement par exemple) sur la 
diversité du zoobenthos (Furse et al., 1984). 

Dans cette étude, la diversité des communautes d'invertébrés est estimée a partir 
de huit descripteurs: (1) le nombre total de taxons (genre ou famille), (2) le nombre de 
taxons sensibles aux pollutions (plécopteres, heptagéniidés et trichopteres a fourreau), 
(3) le rapport nombre de taxons sensibles sur nombre total de taxons (2/1 en %), (4) 
l'indice RIVAUD calculé à partir de I et 2 (Tab 1), (5) le nombre de familles, (6) le 
nombre de familles sensibles aux pollutions (plécopteres, heptagéniidés et trichopteres 
a fourreau), (7) le rapport nombre de familles sensibles sur nombre total de familles 
(6/5 en %), (8) l'indice de diversité de Margalef (Magurran, 1988) dont l'expression est: 


Diversité = (nombre de familles -1) / In du nombre d'individus récoltés. 


Les descripteurs retenus ici sont ceux généralement utilisés pour l'étude des 
cours d'eau (Fore er al., 1996). Certains sont très semblables entre eux (1 à 3 et 5 à 7): 
tout ce qui les différencie, c'est le niveau d'identification atteint (le genre ou la famille). 
Leur inclusion permet d'évaluer l’effet de ce facteur sur l’analyse des résultats. 

Les diversités calculées ici ne sont pas basées sur les espèces mais sur des 
genres, des familles et même des classes (Annexe 1). Cependant cette approche sim- 
plifiée peut être utilisée parce qu'il existe une corrélation très significative entre le 
nombre de familles et d'espèces présentes dans les rivières (Wright et al., 1998). De 
plus, l'analyse des communautés d’invertebres benthiques donne souvent des résultats 
analogues qu'elle soit basée sur des identifications faites au niveau au niveau de la 
famille ou de l’espèce (Furse et al., 1984). 

Les variations des huit descripteurs sont analysées en fonction de l'altitude des 
165 stations de prélèvements, de la région où celles-ci se trouvent (Jura, Jorat et Alpes) 
et de la date des campagnes de prélèvements. Trois campagnes sont comparées entre 
elles: 1989-1991, 1992-1994, 1995-1997. Quatre zones d'altitude sont définies à partir 
des valeurs quartiles (25 %, 50 %, 75 %) et extrêmes de l’altitude des stations de 
prélèvements. 

En plus de l’analyse basée sur les stations, les valeurs moyennes des huit des- 
cripteurs sont calculées pour chaque rivière et pour chaque campagne de prélèvements 
afin d'évaluer la tendance rivière par rivière. Pour ce faire, la valeur du coefficient de 
corrélation de rang de Spearman est calculée entre les trois valeurs moyennes de chaque 
descripteur et les dates correspondantes des trois campagnes de prélèvements. Le 
nombre de rivières où la valeur du descripteur augmente régulièrement (r, = 1.00) est 
ensuite comparée à celui des rivières où sa valeur diminue (rs = - 1.00) au moyen du 
test binomial. Les rivières où les valeurs de r, sont autres que +1 ou -1 sont exclues de 
la comparaison. 


12 CLAUDE LANG 


RESULTATS 


Le tableau 2 présente l’évolution des valeurs moyennes des huit descripteurs du 
zoobenthos en fonction de trois critères: |’ altitude, la région et la date des campagnes de 
prélèvements. Nous constatons tout d'abord que les valeurs moyennes des huit 
descripteurs augmentent significativement avec l’altitude des stations de prélèvement. 
C'est entre 375 m et 449 m d'altitude, dans la zone où les impacts des activités 
humaines sont les plus forts (Lang & Reymond, 1995), que les valeurs observées sont 
les plus basses. Nous observons ensuite que, dans les rivières des Préalpes et des Alpes, 
les nombres moyens des taxons et des familles sensibles aux pollutions sont nettement 
plus élevés dans que dans celles du Jura et du Jorat. Comme la diversité totale est la 
même dans les trois régions, le pourcentage de taxons et de familles sensibles aux 
pollutions est plus élevé dans les rivières de montagne qu'ailleurs. Dans l'interprétation 


TABLEAU 2. Valeurs moyennes par station de 8 descripteurs du zoobenthos en fonction de l'alti- 
tude, de la région et de la campagne de prélèvements. Descripteurs: nombre total de taxons 
d'invertébrés (NT), nombre de taxons sensibles aux pollutions (NTS), pourcentage de taxons 
sensibles aux pollutions (PNTS), indice RIVAUD, nombre de familles (NF), nombre de familles 
sensibles aux pollutions (NFS), pourcentage de familles sensibles aux pollutions (PNFS), indice 
de diversité de Margalef (DIV). Au sein d'un même ensemble et d'une même variable, les 
moyennes qui ne sont pas significativement différentes au seuil de 5 %, sont indiquées par des 
lettres semblables. Ensembles 1 et 2 basés sur tous les résultats 1989 - 1997. 


Ensembles Valeurs moyennes par station 

comparés N° NT NTS PNTS RIVAUD NF NES PNFS DIV 

1. Altitude (m) 375-449 43 18.9A 36A 172A 84A 174A 2.9A 15.6A 2.16A 
450-539 43 241B 7.0B 27.9B 12.3B 20.9B 5.2B 244B 252B 


340-699 48 253B 71.867 292B7 12.9B 21.755.6BE252BE2>7B 
700-1410 41 26:18 104€ 40:9C 14.7C7722.0B TIC ACC 


2. Région Jura) 15 7229A 60A 237A ILIA 20NA 45k 210A 
Jorat®? 47 243A 64A 24.2A 11.8A 21.2A 4.8A 214A 2.53A 
Alpes? 43 23.7A 10.0B 42.2B 13.9B 20.2A 6.7B 34.0B 2.62A 


3. Campagne 1989-1991 165 21.7A 64A 28.0A 11.1A 19.0A 48A 244A 238A 
1992-1994 165 226A 7.0A 29.1A 11.8A 19.7A 5.1A 247A 246A 
1995-1997 165 26.2B 8.0B 28.9A 13.2B 22.6B 5.7B 24.4A. 2.68B 


a) Nombre de stations 

b) Altitude des stations: 375 - 1040 m, moyenne: 522 m 
©) Altitude des stations: 430 - 775 m, moyenne: 555 m 
d) Altitude des stations: 380 - 1410 m, moyenne: 882 m 


des résultats, il faut noter que les effets de l’altitude et de la région sont souvent meles. 
Par exemple, les valeurs moyennes observées entre 700 m et 1410 m d’altitude corres- 
pondent en majeure partie aux rivières des Alpes et des Préalpes: les valeurs observées 
en-dessous de 450 m, aux rivieres du Jura. Remarquons enfin que les valeurs moyennes 
de six des huit descripteurs du zoobenthos augmentent entre 1989 et 1997. C'est entre la 
deuxieme et la troisieme campagne de prelevements que l’augmentation est parti- 
culièrement marquée. 


DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 113 


22- 
20 - © Press 
18- == 
©) 
164 == 
(tl el 
14 = 
na] — 
a 
5 12, tet) 
<x 
> 
G 10a 
SES] 
8- | — © O 
— @ 
6- ©) O 
4, = ©) -— 
x 
27 = = 
0 T T 
N= 43 43 38 41 43 43 38 41 43 43 i 38 41 
1989-1991 1992-1994 1995-1997 
CAMPAGNES 
FIG. 2 


Variations des valeurs quartiles (petits côtés du rectangle de bas en haut: 25%, 50%, 75%) et 
extremes (lignes) de l'indice RIVAUD en fonction de l’altitude des stations de prélèvements au 
cours des trois campagnes effectuées entre 1989 et 1997. Les cercles et les étoiles indiquent des 
valeurs aberrantes excédant 1.5 fois l’espace interquartile. Pour chaque campagne, les stations 
sont réparties en 4 classes d’altitude de gauche à droite: 375 - 449 m, 450 - 539 m, 540 - 669 m, 
700 - 1410 m. N = nombre de stations par classe d’altitude. 


Les figures 2 et 3 montrent d’une part que les valeurs quartiles de l'indice 
RIVAUD et du nombre de familles augmentent régulièrement avec l’altitude des sta- 
tions dans deux des trois campagnes de prélèvements comparées, d'autre part que les 
différences entre les trois campagnes sont également significatives (analyse de 
variance: effet de l’altitude, P = 0.001; effets de la campagne, P = 0.001). Remarquons 
cependant que, pour la campagne 1989 - 1991, les valeurs médianes de la diversité sont 
les mémes dans les trois dernières zones d'altitude. Cette répartition particuliere semble 
indiquer que la sécheresse exceptionnelle de 1989 a exercé une influence negative sur la 
diversité du zoobenthos (Fig. 4). 

La corrélation positive (r, = 0.82) qui existe entre le nombre de taxons sensibles 
aux pollutions et le nombre total de taxons présents dans les 47 rivières vaudoises 
étudiées entre 1989 et 1997 (Fig. 5), permet de classer celles-ci en fonction de la 


114 CLAUDE LANG 


40, 
35 - 
(O) 

30 
v 
Lu 
Si) 
Er] 
> 4 
= 25 
La 
LUI 
(cn —_—— 
Lu 20 + 
œ 
faa) | 
= | Pa] | 
oO | | ar: 
z= — 

154 

| a o 
10, O 
© 
5 T T T 
N= 43 43 38 41 43 43 38 41 43 43 38 41 
1989-1991 1992-1994 1995-1997 
CAMPAGNES 
FIG. 3 


Variations des valeurs quartiles et extrêmes du nombre de familles d'invertébrés en fonction de 
l'altitude des stations de prélèvements au cours des trois campagnes effectuées entre 1989 et 1997 
(légende, voir Fig. 2). 


diversité du zoobenthos. En bas et à gauche de la figure, nous trouvons de petites 
rivières, situées en général à basse altitude, dans des paysages très modifiés par 
l’homme: la Dullive, la Petite Glâne, la Morges et le Boiron de Morges. En haut a 
droite, nous observons des rivières de montagne comme l’Hongrin et la Torneresse qui 
drainent des bassins versants relativement peu modifiés par l’homme, tout au moins en 
amont des stations étudiées. Certaines rivières de montagne, notamment l’Avancon, la 
Tinière, la Veveyse et la Grande Eau, occupent une position marginale sur la figure 5. 
La diversité des taxons sensibles aux pollutions (surtout celle des plécoptères) y est très 
élevée par rapport à la diversité totale. Cette situation particulière s'explique par un 
caractère torrentiel accentué qui élimine de nombreux taxons. Comparons enfin la 
faible diversité observée dans la Venoge à la diversité élevée de son principal affluent 
le Veyron. Si la qualité de l'environnement s’améliorait à basse altitude, la situation de 
la Venoge devrait se rapprocher de celle du Veyron. 

L'évolution de la diversité du zoobenthos entre 1989 et 1997, pour les 47 
rivières étudiées. est présentée dans le tableau 3 avec les valeurs moyennes des huit 
descripteurs utilisés. Sur les 376 tendances calculées (8 descripteurs multipliés par 47 


DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 115 


TABLEAU 3. Valeurs moyennes et tendances de 8 descripteurs du zoobenthos calculées pour 
chacune des 47 rivieres vaudoises visitées a 3 reprises entre 1989 et 1997. Tendance: augmen- 
tation (+), diminution (-), aucune tendance (0). Descripteurs: nombre total de taxons (NT), 
nombre de taxons sensibles aux pollutions (NTS), pourcentages des taxons sensibles aux 
pollutions (PNTS), indice RIVAUD, nombre total de familles (NF), nombre de familles sen- 
sibles aux pollutions (NFS), pourcentage des familles sensibles aux pollutions (PNFS), indice 
de diversité de Margalef (DIV). 


Riviere Valeurs moyennes (en dessus) et tendances (en dessous) 
No (Stations) NT NTS  PNTS RIVAUD NF NFS = PNES DIV 
1 Doye 15.3 1.7 11 DA 15.3 1.7 11 1.9 
(1) 0 0 0 0 0 0 0 0 
2 B. de Nyon 1557 1.8 10.6 6 15.1 1.8 11 1.9 
(4) 0 0 0 0 0 0 0 0 
3 Asse 19.4 3.8 16.5 8.3 17.8 3.] 15.6 DE 
(4) + 0 0 + + 0 0 + 
4 Promenthouse 23.8 6.7 DAD. 12 2A 4.9 22.8 2.6 
(3) + + + + + + + 0 
5 Colline 255 WS 28.3 13 21.8 5.5 24.7 DT] 
(2) 0 + + 0 0 + + 0 
6 Cordex 26.3 9.7 36.6 14.7 23.3 TI 30 2.8 
@ + + + 0 0 0 0 - 
7. Serine 20 6.2 30.6 11 18.7 4.8 26.3 DES 
(2) 0 0 - 0 0 0 0 0 
8 Dullive 10.7 0.5 I 3.5 10.7 0.5 I 1.3 
(2) 0 0 0 + 0 0 0 0 
9 Aubonne DD 9.3 33.9 14.5 22.9 6.2 25) 297: 
(5) + 0 0 + + + 0 0 
10 Toleure 26.8 10.3 38.6 14.8 22.8 Ue) 33 2.9 
(2) 0 + + 0 0 0 0 - 
11 Saubrette 18.3 4.7 25.3 10 LZ 4 22.9 2.1 
(1) 0 0 0 0 0 0 0 0 
12 B. Morges 18 2.6 13.2 ES 17.2 DIS 13.4 DAI 
(5) 0 0 + 0 0 0 + 0 
13 Morges 18 3 16.1 8 17.1 2.9 16.5 2 
(5) 0 - - 0 0 0 0 0 
14 Venoge 213 4.5 195 9.8 19 35 WZ 2.4 
(14) + 0 0 + + 0 0 + 
15 Veyron 30.9 10.2 32.9 15.8 24.8 TS) 29.2 5 
(4) + + 0 + + 0 0 + 
16 Nozon 28.9 10.8 37.1 1537 23.4 7.4 31.6 DET 
(6) 0 + 0 0 0 + 0 0 
17 Orbe DES 6.3 22.9 125 233 Sl DIET 2.8 
(5) + + + + + + + + 
18 Mujon 17.3 2 11.4 7 17.3 2. 11.4 1.9 
(1) 0 0 - 0 0 0 = È 
19 Arnon 30.9 10.2 32.9 15.8 24.8 18 29.2 3 
(7) 0 0 + + 0 0 + 0 
20 Baumine 28.7 Dall 19.3 13 25 4.7 18.3 2.8 
(1) 0 0 0 0 0 0 (0) 0 
21 Talent 19 4.] 18.4 8.6 11927 3.4 17.3 22 
(7) + + + + + + + + 
22 Buron DS 6.3 2/61 12 20.7 4.7 22.8 DIS 
(2) 0 0 0 0 0 0 0 0 
23 Mentue 275 8.5 30 14.1 22.9 5.8 24.5 DI] 
(8) 0 0 0 0 0 0 0 0 


116 CLAUDE LANG 


TABLEAU 3 (suite) 


Rawiere Valeurs moyennes (en dessus) et tendances (en dessous) 


No (Stations) NT NTS PNTS RIVAUD "NE NFS PNES DIV 


24 Sauteru 29 


7.8 DAN 14.2 26 6.2 24 3). il 
(2) 0 0 + 0 0 0 0 0 
25 Betite:;Gläane 215.8 12 1:3 SI 15.6 122 123 2 
(4) 0 + - + 0 + + 0 

26 Broye 27.4 7 24.4 1331 23.4 53 22 2.8 
(7) + 0 0 + + 0 0 + 

27 Arbogne 23.9 5 20.4 11 21 5 18.1 DIS 
(3) + + + + 0 + + 0 

28 Lembe 15.7 DEN 16.5 MI 138 2.7 16.9 1.9 
(1) 0 - 0 - 0 - - 0 

29 Cerjaule DS 8.3 32.4 197 23 N 30.5 2.8 
(1) 0 - 0 - 0 - 0 - 

30 Mérine 313 EY, 37.1 16.7 IST 8 312 SA 
(1) 0 0 0 0 0 0 - 0 

31 Bressonne 27.8 93 32.6 14.4 23.6 6.7 28 2.9 
(3) + + + + 0 + + + 

32 Carrouge 21.3 4.8 21.4 10.2 19.5 4 20 22 
(2) 0 0 0 0 0 0 0 - 

33 Grenet 26.4 8.2 30.8 13.8 22.8 62 ZII ZA) 
(4) 0 0 0 0 0 0 - + 

34 Mionne 2957 10.3 34.7 1527 24.7 TR SISI DS) 
(1) 0 0 0 0 0 0 + 0 

35 Forestay 223 3 12.7 9 20.3 3 14.1 2.4 
(1) 0 0 0 0 0 0 0 0 

36 Veveyse DI. 8.7 39.3 13 18 5 doll 2.4 
(1) + 0 0 + u 0 0 + 

37 B.Clarens 29.4 [EZ 37.9 15.8 24.8 7.6 30.4 DIO, 
(4) 0 0 0 0 + 0 0 + 

38 B. Montreux 27.6 IH 40.3 1577 23.6 TE? 30.6 DIO, 
(3) 0 0 0 0 0 0 - + 

39 Tinière 19.7 7.9 38.8 11.6 17.4 SLI 30 DS 
(3) 0 0 0 0 (0) 0 0 0 

40 Grande Eau 20.8 9.5 45.7 13 17:9 6.7 STES) 2.4 
(10) + - 0 0 0 0 - 0 

41 Gryonne 23.8 8.9 37.6 13.6 20.8 6.1 29.6 2 
(4) 0 0 - 0 0 - - + 

42 P. Gryonne 25 8 32 153 225 SIT 252 ei 
(1) 0 + + (0) 0 0 + + 

43 Avancon 17.4 TS) 45.4 11.2 15.2 SA ST DED 
(7) + + - + 0 + - + 

44 Hongrin 327 1235 38.1 1783 26.7 8 30 3,2 
(2) + + 0 + + + 0 + 

45 Sarine 26.8 Fi] 42.2 152 223 TD 34.6 2.8 
(4) + 0 0 + + 0 0 + 
46 Flendruz 28 123 44.] 16.3 28 8 3922 3 
(1) + + 0 + + 0 - + 

47 Torneresse 30 14.1 47 17.3 24.1 8.6 355 DIO 
(3) 0 0 0 0 0 0 0 * 

Tendance (+/-) (17/0) (16/3) (13/5) (18/2) (3/0) Cais) (11/9) (18/5) 

0 


Probabilite 0 0.004 0.096 0 0.057 0.824 0.012 


DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES AG 


PHOSPHORE 


90 - 


50 - 


40- 


30 + 


20 | T 


SU T T TILE T T = 
1983 1985 1987 1989 1991 1993 1995 1997 
1984 1986 1988 1990 1992 1994 1996 — 1998 


T = T Ter 


ANNEE 


FIG. 4 


Evolution des concentrations moyennes en phosphore soluble (mg / m?) et des débits d'eau (mò / 
sec multiplies par 10) dans la Venoge entre 1984 et 1997. Les concentrations sont pondérées par 
le débit (Orand er al., 1998). 


rivières), 117 indiquent une augmentation de la valeur du descripteur et 27 une 
diminution (test binomial, P = 0.001); dans 232 cas, aucune tendance bien marquée 
n'est observable. Lorsque les tendances sont analysées descripteur par descripteur 
(colonne par colonne), les nombres d'augmentations sont significativement supérieurs à 
ceux des diminutions pour six descripteurs sur huit. Pour les pourcentages de taxons ou 
de familles sensibles aux pollutions, les différences ne sont pas significatives, ce qui 
suggère que ces deux descripteurs sont moins efficaces que les six autres pour détecter 
une tendance. Lorsqu'on examine l’évolution de la diversité rivière par rivière (ligne 
par ligne), la tendance dominante est clairement visible dans la plupart des cas, même si 
l'ensemble des huit descripteurs n'indiquent le même diagnostic que dans dix rivières. 
Les diagnostics contradictoires (une augmentation et une diminution dans la même 
rivière) sont également rares: ils sont le fait de l'indice de Margalef et du pourcentage 
de taxons ou de familles sensibles aux pollutions, descripteurs dont la valeur indicatrice 
a déjà été mise en doute ci-dessus. 


118 CLAUDE LANG 


167 | 


14, o 
124 


105 40 


TAXONS SENSIBLES 
à 


2- EG 12 a 


10 15 20 25 30 35 


TOUS LES TAXONS 


Fic. 5 


Relation entre le nombre total de taxons d'invertébrés et le nombre de taxons sensibles aux 
pollutions (plécopteres, heptagéniidés et trichopteres à fourreau), permettant de classer les 47 
rivieres vaudoises visitées entre 1989 et 1997, les unes par rapport aux autres en fonction de leur 
diversité (voir Tab. 3, numéros d'identification et noms des rivieres). 


DISCUSSION 


Dans les rivières vaudoises, la diversité du zoobenthos augmente d’une part en 
fonction de l’altitude des stations de prélèvements dans chacune des trois campagnes 
effectuées entre 1989 et 1997, d’autre part en fonction du temps écoulé entre la 
première et la troisième campagnes (Figs 2, 3). La première tendance correspond à 
l'augmentation amont-aval des perturbations d’origine humaine qui s’observe dans la 
plupart des bassins versants vaudois (Lang & Reymond 1993, 1995). La deuxième 
tendance montre à la fois les progrès de l'épuration des eaux et les effets d'une météo- 
rologie favorable. 

L’épuration des eaux a permis de contrôler efficacement certains polluants ainsi 
que l’illustre la baisse significative (r, = - 0.84, n = 14) des concentrations en phosphore 
soluble dans la Venoge entre 1984 et 1997 (Fig. 4). Cette figure montre également que 
la longue sécheresse de 1989 a momentanément compromis l’amélioration de la qualité 


DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 119 


chimique de l’eau. En effet cette année-la, le debit moyen de la Venoge est 
particulierement faible, comme d’ailleurs celui des autres rivieres vaudoises. A cause de 
ce manque d’eau, les apports en phosphore soluble, ainsi que ceux d'autres polluants, 
sont moins dilués et les concentrations plus élevées qui en résultent, affectent davantage 
le zoobenthos (Allan, 1995). 

La situation météorologique extreme de 1989 a influence fortement la premiere 
des trois campagnes de prélèvements effectuées entre 1989 et 1997 (Figs 2, 3). 
L'évolution ultérieure de la diversité pourrait ainsi s’interpreter comme le rétablisse- 
ment progressif du zoobenthos après un épisode climatique exceptionnel. Si cette 
interpretation est correcte, la diversité du zoobenthos observée pendant la dernière cam- 
pagne correspondrait à une situation normale sur le plan météorologique pour les 
rivières vaudoises. En prenant les diversités observées en altitude comme valeurs de 
référence pour un milieu où les perturbations d’origine humaine sont minimes (Figs 2, 
3), nous constatons une sérieuse baisse de la diversité à basse altitude, ceci malgré une 
épuration des eaux complètement réalisée (Fiaux ef al., 1998). 

L'effet de l'amélioration de la qualité chimique de l’eau sur la diversité du 
zoobenthos serait ainsi contrecarré par la persistance d'autres perturbations d’origine 
humaine (Allan, 1995). Citons entre autres: l’alluvionnement, l’altération des débits de 
crue et d’étiage et la dégradation de la végétation riveraine. En un mot, tout ce qui 
modifie le caractère naturel d'un cours d'eau et de son bassin versant fait diminuer la 
diversité du zoobenthos. Le milieu naturel environnant les rivières vaudoises est 
dégradé mais l'étendue de sa dégradation ne peut pas être évaluée, faute d'une base de 
référence et de données quantitatives précises. Cependant, l'influence de | agriculture et 
de l'urbanisation est évidente. 

Les activités agricoles pratiquées dans le bassin versant influencent la diversité 
du zoobenthos. L'évolution d'une rivière américaine illustre ce point (Grubaugh & 
Wallace, 1995): après la disparition dans le bassin versant d'une agriculture intensive 
(utilisant beaucoup d'engrais, de pesticides et provoquant une forte érosion), la 
diversité du zoobenthos a fortement augmenté. Dans le canton de Vaud, l’évolution 
inverse s’est produite entre 1960 et 1970 avec le passage d’une agriculture tradi- 
tionnelle à une agriculture intensive (B. Reymond, com. pers.). Il est significatif de 
constater que la diversité des plécoptères, insectes particulièrement sensibles à l’in- 
fluence humaine (Fore er al., 1996), a fortement diminué entre 1945 et 1982 dans le 
cours inférieur de la Venoge, du Talent, de la Broye et d’autres cours d’eau traversant 
des paysages agricoles très modifiés par l’homme (Aubert, 1984). 

En plus des effets à long terme de l’agriculture sur le paysage, des pollutions 
tant aiguës que chroniques par des produits phytosanitaires pourraient expliquer la 
faible diversité du zoobenthos qui persiste entre 1989 et 1997 dans certaines rivières 
vaudoises où l’épuration des eaux est entièrement réalisée. Citons, par exemple, le 
Boiron de Nyon, l’Asse, la Dullive, le Boiron de Morges et la Morges (Tab. 3). En 
raison de la conjonction de faibles débits et d'une agriculture très active, divers produits 
phytosanitaires sont présents dans l’eau en concentrations relativement élevées (Corvi 
& Kim-Heang, 1997: Vioget & Strawczynski, 1997). De plus, des déversements acci- 


120 CLAUDE LANG 


dentels de ces produits provoquent frequemment la mort des poissons et des inver- 
tebres, parfois seulement celle des invertebres; dans ce cas, les pollutions et leurs 
auteurs sont particulierement difficiles a detecter (Ph. Tavel, com. pers.). 

En plus de ces pollutions aigués mais sporadiques, les concentrations en phyto- 
sanitaires mesurées dans ces cours d'eau suggèrent qu'il pourrait exister une pollution 
chronique. Celle-ci agirait de facon subtile sur le taux de survie des especes. Par 
exemple, le nombre de trichoptères du genre Limnephilus, capables d’émerger de l’eau 
et donc de se reproduire, diminue significativement en présence d'une tres faible 
concentration (1 ng/l) de lindane, un insecticide (Schulz & Liess, 1995). Il est bien clair 
que des effets si subtils ne peuvent pas étre mis en évidence par des méthodes clas- 
siques de surveillance biologique des eaux. 

Le zoobenthos des rivieres vaudoises subit également les effets négatifs d’une 
urbanisation accrue, surtout a basse altitude. En effet, la densité de la population 
humaine qui constitue un bon indicateur des impacts lies a urbanisation (Statzner & 
Sperling, 1993), augmente en général d’amont en aval dans les bassins versants du 
canton de Vaud (Lang & Reymond, 1995). Une forte densité humaine entraine par 
exemple: (1) l'augmentation des surfaces de sol imperméables ce qui modifie les débits 
de crue et d’étiage, facteurs critiques pour le zoobenthos; (2) l’accroissement du réseau 
de canalisations qui conduisent les eaux usées aux stations d’épuration (Statzner & 
Sperling, 1993). Plus ce réseau est étendu, plus la matiere organique et les polluants 
peuvent s’y accumuler par temps sec: de ce fait, plus grand sera impact de ces dépôts 
sur la riviere apres une forte pluie, sauf si les déversoirs d’orage sont congus et entre- 
tenus de facon optimale. 

En conclusion, Statzner & Sperling (1993) suggèrent qu'une rivière épurée, son 
bassin versant et la population qui l'habite constituent un système complexe qu'il 
faudrait gérer comme un tout, en utilisant différentes approches, si l'on veut améliorer à 
moindre coût la qualité biologique de l’eau. C’est une conception analogue qui devrait 
être adoptée à l’avenir dans les rivières vaudoises. 


REMERCIEMENTS 


La collaboration d'Olivier Reymond tant sur le terrain qu’en laboratoire m’a 
permis de mener à bien cette étude. Les commentaires de Michel Dethier, de Philippe 
Gmür, d’Andres Strawczynski et de Philippe Vioget ont contribué à améliorer cet 
article. 


BIBLIOGRAPHIE 


ALLAN, J. D. 1995. Stream ecology. Chapman et Hall, London, 388 pp. 

AUBERT, J. 1984. L’atlas des Plécoptères de Suisse - Influence de la pollution. Annales de 
Limnologie 20: 17-20. 

CORVI, C. & KHIM-HEANG, S. 1997. Surveillance des produits phytosanitaires dans les eaux des 
affluents du bassin lémanique, campagne 1995 - 1996. Rapport de la commission inter- 
nationale pour la protection des eaux du Léman contre la pollution: 125-144. 

FIAUX, J.-J.. VALLIER, R. & VIOGET, PH. 1998. Stations d'épuration, bilans 1997. Service des eaux 
et de la protection de l'environnement. CH-1066 Epalinges. 


DIVERSITE DU ZOOBENTHOS DANS 47 RIVIERES 12] 


Fore, L. S., KARR, J. R. & WISSEMAN, R. W. 1996. Assessing invertebrate response to human 
activities: evaluating alternative approaches. Journal of the North American Bentho- 
logical Society 15: 212-231. 

FURSE, M. T., Moss, D., WRIGHT, J. F. & ARMITAGE, P. D. 1984. The influence of seasonal and 
taxonomic factors on the ordination and classification of running-water sites in Great 
Britain and on the prediction of their macro-invertebrate communities. Freshwater 
Biology 14: 257-280. 

GRUBAUGH, J. W. & WALLACE, J. B. 1995. Functional structure and production of benthic 
community in a Piedmont river: 1956-1957 and 1991-1992. Limnology and Oceano- 
graphy 40: 490-501. 

HELLAWELL, J. M. 1986. Biological indicators of freshwater pollution and environmental mana- 
gement. Elsevier Applied Science Publisher, London and New York, 546 pp. 

Hynes H. N. B. 1975. The stream and its valley. Verhandlungen der internationale Vereinigung 
fiir theoretische und angewandte Limnologie 19: 1-15. 

LANG, C. 1996. Qualité de l'environnement indiquée par la diversité du zoobenthos dans les 
rivieres de montagne: campagnes 1985 - 1995. Bulletin de la Société Vaudoise des 
Sciences Naturelles 84: 125-137. 

LANG, C. 1997. Qualité biologique de 37 rivières vaudoises en 1996 indiquée par la diversité du 
zoobenthos. Bulletin de la Société Vaudoise des Sciences Naturelles 84: 323-332. 

LANG, C. 1998. Qualité biologique de 29 rivieres vaudoises en 1997 indiquée par la diversité du 
zoobenthos. Bulletin de la Société Vaudoise des Sciences Naturelles 86: 61-71. 

LANG, C. & REYMOND, O. 1993. Empirical relationships between diversity of invertebrate 
communities and altitude in rivers: application to biomonitoring. Aquatic Sciences 55: 
188-196. 

LANG, C. & REYMOND, O. 1995. An improved index of environmental quality for Swiss rivers 
based on benthic invertebrates. Aguatic Sciences 57: 172-180. 


MAGURRAN, A. E. 1988. Ecological diversity and its measurement. Croom Helm, London and 
Sydney, 179 pp. 

ORAND, A., DORIOZ, J.-M. & GAGNAIRE, J. 1998. Bilan des apports par les affluents au Léman et 
au Rhone à l'aval de Genève. Rapport de la commission internationale pour la pro- 
tection des eaux du Léman contre la pollution: 117-133. 


RIBAUT, J. P. 1966. Les poissons des cours d'eau du canton de Vaud (Suisse). Mémoire de la 
Société Vaudoise des Sciences Naturelles 87: 69-128. 

REYMOND, O. 1995. Surveillance biologique des cours d’eau: matériel et méthode pour trier les 
prélevements d'invertébrés. Bulletin de la Société Vaudoise des Sciences Naturelles 83: 
209-215. 

SCHULZ, R. & LiEss, M. 1995. Chronic effects of low insecticide concentrations on freshwater 
caddisfly larvae. Hydrobiologia 299: 103-113. 

STATZNER, B. & SPERLING, F. 1993. Potential contribution of system specific knowledge (SSK) to 
stream management decision: ecological and economic aspects. Freshwater Biology 29: 
313-342. 

TACHET, H., BOURNAUD, M. & RICHOUX, PH. 1980. Introduction à l'étude des macroinvertébrés 
des eaux douces. Université de Lyon, 156 pp. 

VIOGET, PH. & STRAWCZYNSKI, A. 1997. Phytosanitaires dans les cours d'eau vaudois du bassin 
du Rhin et du Rhône. Notes du service des eaux et de la protection de l'environnement, 
CH-1066 Epalinges. 

WRIGHT, J. F., Moss, D. & FURSE, M. T. 1998. Macroinvertebrate richness at running-water sites 
in Great Britain: a comparison of species and family richness. Verhandlungen der inter- 
nationale Vereinigung fiir theoretische and angewandte Limnologie 26: 1174- 1178. 


122 


CLAUDE LANG 


ANNEXE |. Fréquence des taxons identifiés entre 1989 et 1997. Source: Lang 1996, 1997, 1998. Taxons 
présents +, 1 - 10% des prélèvements, fréquents ++, 11 - 50%, très fréquents +++, 51 - 100%, - absent. 


L'astérisque signale les taxons sensibles aux pollutions. 


Région 
Taxons®) Jura Jorat Alpes 
TURBELLARIA 
Planariidae 
Polycelis ++ + + 
Dugesiidae 
Dugesia ++ ++ ++ 
Oligochaeta +++ +++ ar 
HIRUDINEA 
Glossiphonidae 
Helobdella + + & 
Glossiphonia + aft 
Erpobdellidae 
Erpobdella ++ ++ + 
MOLLUSCA 
Hydrobiidae + = + 
Ancylidae ++ ++ x 
Limnaeidae ++ + At, 
Sphaeriidae ++ ++ + 
Physidae + = 3 
Bythinellidae + = 4 
Bithyniidae = = + 
HYDRACARINA +++ ++ ++ 
CRUSTACEA 
Gammaridae +++ +++ ++ 
Asellidae + + + 
EPHEMEROPTERA 
Ephemeridae 
Ephemera + + En 
Heptageniidae 
Epeorus* +++ +++ ++ 
Rithrogena* +++ ae +++ 
Ecdyonurus* ++ +++ +++ 
Heptagenia* = = n 
Caenidae 
Caenis + sus si 
Baetidae +++ +++ +++ 
Ephemerellidae 
Ephemerella + SEE n 
Torleya - + + 
Leptophlebiidae 
Habrophlebia ++ ++ CE 
Habroleptoides ++ ++ ++ 
Paraleptophlebia + + + 
PLECOPTERA 
Taenioptery gidae 
Taeniopteryx* - < = 
Brachyptera’ ++ ++ ++ 
Rhabdiopteryx* - + 44 
Nemouridae 
Amphinemura* ++ -- a 
Protonemura* ++ ++ +++ 
Nemoura* ++ ++ ae 
Leuctridae 
Leuctra* +++ ++ +++ 
Capniidae 
Capnia* + + + 
Capnioneura* = z + 
Chloroperlidae 
Chloroperla* + = ++ 
Perlodidae 
Isoperla* ++ en sen 
Perlodes* - = + 
Perlodidae gen.* - = at 


a) Identifiés d'après Tachet er al. (1980). _ 


Sciomyzidae 


Région 
Taxon Jura Jorat Alpes 
Perlidae 
Perla* + È ann 
Dinocras* - = fi 
ODONATA - + = 
HETEROPTERA - = n 
PLANIPENNIA 
Osmylidae 
Osmylus # È + 
MEGALOPTERA 
Sialidae 
Sialis + AL + 
COLEOPTERA 
Gyrinidae + + È 
Dytiscidae + pù ni 
Haliplidae ch + + 
Hydrophilidae + + + 
Helodidae + + A 
Eubridae = + a, 
Dryopididae - + & 
Hydraenidae 
Hydraena ++ ++ ++ 
Ochthebius = A a 
Elmidae 
Elmis ++ EME run. 
Esolus ++ + + 
Limnius tato +++ ++ 
Normandia + = È 
Riolus ++ ++ + 
Oulimnius + + + 
Dupophilus = = A 
TRICHOPTERA 
Rhyacophilidae +++ +++ +++ 
Glossosomatidae * a + er 
Hydroptilidae* ++ ++ + 
Hydropsychidae +++ +++ ++ 
Philopotamidae - + sit 
Polycentropidae + + + 
Psychomiidae +++ ++ + 
Phryganeidae* - = 2 
Limnephilidae* +++ +++ ++ 
Drusinae* = = i 
Goeridae* + + AL 
Beraeidae* + 2 + 
Odontoceridae* ++ ++ a 
Sericostomatidae* Di ++ Er 
Lepidostomatidae* + + Be 
Leptoceridae* + = È 
DIPTERA 
Blephariceridae + + ++ 
Tipulidae ++ - Sto 
Limonidae ++ +++ +++ 
Psychodidae ++ ++ ++ 
Simuliidae +++ +++ +++ 
Chironomidae +++ +++ it 
Ceratopogonidae ++ +++ ++ 
Stratiomyidae + + + 
Empididae +++ +++ ++ 
Dolicopodidae + + + 
Athericidae ++ ++ +++ 
Dixidae = + + 
Anthomyidae ~ - + 
Tabanidae + + È 

+ = = 


REVUE SUISSE DE ZOOLOGIE, 107 (1): 123-138; mars 2000 


Contribution a la connaissance de la famille Eucinetidae 
(Coleoptera) 


Stanislav VIT 
Rue de la Poterie 26, CH-1202 Geneve, Suisse. 


Contribution to the knowledge of Eucinetidae (Coleoptera). - A new 
genus, Proeuzkus gen. n., is described to accomodate two new species, 
P. pachys from Nepal and P. coecus from Thailand. Proeuzkus pachys 
sp. n. (type-species) represents the first record of a highly specialised form 
of Eucinetidae from the area extending from the Caucasus to Japan, while 
P. coecus sp. n. is the first anophthalmous species of the family. Another 
morphologically interesting species Tohlezkus laticanthus Sp. n. is descri- 
bed from Malaysia, and Eucinetus xaca sp. n. with a highly derived aedea- 
gus is added to the Mexican fauna. 


Key-words: Coleoptera - Eucinetidae - Proeuzkus gen. n. - Tohlezkus - 
Eucinetus - taxonomy - new species - Nepal - Thailand - Malaysıa - 
Mexico. 


INTRODUCTION 


La famille des Eucinetidae comporte actuellement six genres et 38 especes 
valides, dont 28 (plus deux sous-especes) dans le seul genre Eucinetus Germar, 1818. 
Les cinq genres restants refletent une grande hétérogénéité de combinaisons des 
caracteres morphologiques. Les Eucinetidae font partie des six lignées phyletiques de 
Coléopteres aux comportements alimentaires corrélés avec des spores. Plusieurs 
especes (larves comprises) sont connues pour étre associées a divers champignons et ıl 
reste admis que certaines d’entre elles sont associées à des Myxomycetes (Lawrence & 
Newton, 1980; Chandler, 1991). Les premieres formes a morphologie buccale haute- 
ment spécialisée ont été décrites dans les genres Tohlezkus Vit et Jentozkus Vit (Vit, 
1977), puis l’ensemble des données progressivement revu et remanié (Vit, 1981, 1985, 
1990, 1995). Aucune hypothese sur les relations phylogénétiques au sein de la famille 
n’a encore été publiée. En l’état de nos connaissances, on y soupçonne cependant une 
grande fréquence des caractères convergeants. 

Parmi des matériaux inédits, de provenances fort diverses, souvent modestes 
quant au nombre d'individus disponibles, les quatre espèces réunies dans ce travail 
représentent des espèces porteuses de caractères intéressants ou très particuliers. Elles 
me semblent succeptibles d'apporter des éléments nouveaux au regard des relations 
phylogénétiques au sein de cette famille, où des caractères vraisemblablement dérivés 
se retrouvent souvent dans des groupes supposés phylétiquement éloignés. 


Manuscrit accepté le 27.07.1999 


124 STANISLAV VIT 


Un genre nouveau, Proeuzkus gen. n. de la région orientale, est érigé pour 
inclure la premiere espece hautement évoluée de la famille Eucinetidae, recensée jus- 
qu'à present des vastes territoires s’etendant entre l’Iran et le Japon. Deux espèces lui 
sont assignées; P. pachys sp. n., remarquable espèce du Népal (espece-type) et le minis- 
cule et anophtalme P. coecus sp. n. de la Thailande. Une curieuse espece Tohlezkus 
laticanthus sp. n. est décrite de Malaisie et finalement une espece du Mexique, Euci- 
netus xaca Sp. n., remarquable par son édéage, complete la liste. 


Abréviations utilisées: 


CSV Collection S. Vit, Genève, Suisse 

CUIC Cornell University, Ithaca, Etats-Unis 

MHNG Museum d’histoire naturelle, Geneve, Suisse 

ZML Museum of Zoology, Lund, Suede 

Dans les descriptions: A.-Antennes; E.- Elytres; F.-Front; Lo.- Longueur; La. - Largeur; 
m. - médian(ne); P.- Pronotum; s. - sutural(e): 


DESCRIPTIONS 
Proeuzkus gen. n. 
Espece-type: Proeuzkus pachys sp. n. 

Corps petit, en dessous de 1,5 mm, trapu, subglobuleux, rappelant celui des 
Sphaerosoma (Sphaerosomatidae); téguments pigmentés, vigoureusement ponctués et 
pubescents. 

Tête (Fig. 1) large, encastrée dans le pronotum jusqu’au niveau des yeux, repliée 
normalement sur la face ventrale du corps; yeux bien développés, latéraux: insertion 
antennaire entièrement cachée au fond d’une profonde cavité, fermée ventralement par 
le bord inférieur explané de la tête, et dorsalement par l'expansion latérale du front; 
suture frontoclypéale fortement développée (cf. Discussion); appareil buccal (non 
étudié en détail) fortement modifié, probabalement du type suceur: labre court, assez 
peu sclérifié (comme chez Bisaya Reitter); palpes maxillaires quatriarticulés; palpes 
labiaux triarticulés, l’article apical subulé. Antennes (Fig. 7) larges, très compactes; 
formées de 11 articles fortement déprimés: scape asymétrique, pédicelle inséré sur sa 
face ventrale (comme chez Jentozkus Vit). Face ventrale de la tête avec de chaque côté 
un pli transversal caréné inexistant dans les autres genres, qui s’étend de la base du 
submentum jusqu'aux tempes, où il forme une protubérence dentiforme saillant latéra- 
lement en arrière des yeux. Pronotum enveloppant la base des élytres; scutellum 
apparent. Elytres non soudés sur la suture, sans traces de stries longitudinales, mais 
pourvus de 10 stries vestigiales; épipleurs entiers, faiblement élargis dans leur portion 
posthumérale puis progressivement atténués, méchanisme de blocage des élytres au 
repos présent. 

Face ventrale (Figs 3-5). Processus prosternal bien développé: angle médian du 
mésosternum très obtus, ne formant pas de processus mésosternal; cavités mésocoxales 
subcontigués; métasternum grand, ses branches latérales bien développées, non 


LA FAMILLE EUCINETIDAE 125 


fusionnees; épisternes metasternaux libres; épisternes et épimères mésosternaux 
complètement fusionnés; extrémités latérales des hanches postérieures découvertes: 
cing sternites apparents chez les deux sexes; méso- et métatibias avec deux éperons 
apicaux chez les deux sexes. Dimorphisme sexuel limité aux seuls épérons apicaux des 
mésotibias. 

Edéage (Figs 9-10) avec le lobe médian non tubuliforme et les paramères 
déplacés tout à fait dorsalement rappelle celui d’ Eucilodes Vit. 

Discussion. Le genre Proeuzkus est défini par son espèce-type P. pachys sp. n. 
Sa tête peu modifiée, ses yeux bien développés et l’appareil buccal du type inter- 
médiaire rappellent Bisaya Reitter du Lencoran et Eucilodes Vit (paléarctiques), mais 
aussi Euscaphurus Casey (néarctique). Le pli transversal lamelliforme de la face ven- 
trale de la tête, supportant ventralement les yeux, et terminé de chaque côté par une 
saillie dentiforme est unique chez les Eucinetidae, tout comme les antennes de ce type 
particulier. L’appartenance de la deuxième espèce, P. coecus Sp. n., à ce genre, est 
cependant plus problématique car la tête a subi d’importantes modifications: atrophie 
des yeux, évolution des pièces buccales vers le type piqueur-suceur et disparition de la 
fosse antennaire (cf. sous coecus). D’autres adaptations, corrélées avec la reduction de 
la taille et une biologie spécialisée, sont celles déjà connues dans d’autres genres, 
comme Tohlezkus Vit ou Jentozkus Vit. Néanmoins, le recouvrement dorsal de l’inser- 
tion antennaire, les antennes dilatées au niveau du funicule et déprimées, le scape 
asymétrique et comprimé, la carene transversale de la tete (ici vestigiale et réduite a ses 
extrémités latérales), le mésosternum court (angle médian non lancéolé) et dépourvu de 
carene médiane, deux éperons apicaux des pattes, constituent les caracteres communs 
aux deux espèces. P. coecus sp. n. est donc accepté ici dans le genre Proeuzkus en tant 
que son espèce derivee. 


Proeuzkus pachys sp. n. Figs 1, 3-11 


Matériel. Holotype 4, étiqueté: NEPAL, Khandbari District, above Tashigaon, 3100 m 
7.1V.1982, A. & Z. Smetana. (MHNG); Paratypes: 26, mêmes données que l’holotype, (1d 
disséqué et conservé en glycérine). (MHNG); 14, 12 idem HT, “Bakan” W of Tashigaon 
3200 m, 5.1V.1982, A. & Z. Smetana. (MHNG, 1% CSV). 

Espece aptere chez les deux sexes, coloration d’un brun foncé rougeatre, pattes a 
peine plus jaunatres; pubescence courte, couchée; téguments non translucides, brillants, 
entierement densément et fortement ponctués. Corps fortement convexe, trapu, acuminé 
apicalement. Longueur 1,40-1,45 mm, largeur 0,97-1,02 mm, située en avant du milieu 
du corps. 

Téte (Fig. 1). Large, faiblement convexe, retrécissant en arriere des yeux et 
convergeant en arriere; tempes doublées ventralement, en arriere des yeux, par l’extre- 
mité latérale dentiforme de la carène ventrale transversale de la tête; largeur de la tête 
(yeux compris) d’un tiers supérieure à sa longueur médiane (labre non compris), cette 
dernière subégale à la largeur du front. Yeux saillants latéralement et parfaitement 
isolés de la face ventrale de la tête, composés d’une vingtaine de grosses ommatidies, 
rebordés en bas par une fine carène subocculaire indistincte en vue dorsale. Côtés 
explanés du front faiblement sinueux au-dessus de l’insertion antennaire, cette dernière 


126 STANISLAV VIT 


Fics 1-2 


Proeuzkus spp. 1, P. pachys sp. n.: tete, vue dorsale; 2, P. coecus sp. n.: tete, vue dorsale. 
Mesures données en mm. 


entierement cachée au fond d’une profonde fosse antennaire entierement fermee; fron- 
toclypéus court, amplement arrondi en avant, les côtés faiblement rebordes; suture 
frontoclypéale fortement développée, formant un canal profond (qui semble en partie 
invaginé), terminé de chaque côté par une fossette circulaire (cf. Discussion). Appareil 
buccal (Fig. 6) du type spécialisé; mandibules et maxilles indistinctes; labre petit, 
ramassé, peu sclérifié; dernier article des palpes maxillaires grand, étiré en pointe, 
l’apex prolongé par une courte épine hyaline; palpes maxillaires triarticulés, très petits, 
dépassant à peine sur le côté. 

Antennes (Fig. 7) compactes, courtes (à peine aussi longues que la moitié de la 
largeur basale du pronotum), fortement déprimées, très larges (seulement trois fois plus 
longues que larges), leur plus grande largeur située au niveau du 8ème segment; articles 


LA FAMILLE EUCINETIDAE 127 


SATA WE 
cr) JE 
1 ANG D = 
ta if 
No 

dì Od | WE 
LG 1 = Î_, no 
IERI 

) Vig N 3 

, 274 4 

STR 


Fics 3-5 


Proeuzkus pachys sp. n. face ventrale du corps: 3, hanches postérieures, méta - et mésosternum 
(pattes omises); 4, prosternum; 5, patte antérieure. (Mesures données en mm). 


128 STANISLAV VIT 


fermement encastres les uns dans les autres, portant de tres longues soies semi-dressees. 
Scape en grande partie visible, ovoide, asymétrique, protubérant a son bord distal et 
cachant partiellement le pédicelle; ce dernier subcylindrique, aussi long que large, 
inséré sur la face ventrale du scape; articles 3-7 du funicule trés courts, fortement 
transverses (au moins 3 fois plus larges que longs); articles 9 et 10 plus robustes, 
transverses, deux fois plus larges que longs, (évoquant une massue mal différenciée); 
segment apical subovoide, plus large que long, surmonté d’une pubescence apicale. 

Pronotum tres grand, occupant un quart de la longueur totale de l’animal. Base 
doublement sinueuse mais peu projetée en arriere; scutellum grand, triangulaire, entie- 
rement visible: côtés finement rebordés; angles postérieurs aigus, enveloppant la base 
des élytres. Ponctuation forte, plus espacée que celle de la tete ou celle des élytres. 

Elytres sub-hémisphériques, aussi longs que larges ensemble; côtés fortement 
arqués; suture élytrale non soudée; strie juxtasuturale présente, raccourcie, marquée tout 
au plus dans la moitié apicale; interstrie juxtasutural renflé dans le tiers apical. 
Ponctuation grossiere et dense, non alignée en stries longitudinales; dix stries ponctuées 
demeurent cependant nettement distinctes sur chaque élytre en milieu aqueux, les stries 
2 et 4 raccourcies, ne dépassant pas le tiers basal. 

Face ventrale (Figs 3-5) brillante et aussi fortement ponctuée que la face dorsale, 
aire médiane faiblement déprimée. Epipleures entiers, faiblement élargis dans leur 
portion posthumérale, puis longuement atténués. Limite médiane du mésosternum 
simplement obtuse, angle apical renflé, glabre, non étiré en un processu médian sépa- 
rant les hanches: métasternum grand, sutures transverses et suture médiane indistinctes; 
languette métasternale bien développée, courte, subparallèle; metépisternes trapus, sub- 
triangulaires (comme chez Eucinetus apterus Vit, 1990), le bord externe explané, 
formant un dispositif de blocage des élytres; mesépimeres transverses, fusionnés avec 
les mesépisternes, la suture indistincte, leur structure tégumentaire cependant diffé- 
renciée. Plaques coxales larges, mais orientées longitudinalement, laissant les extré- 
mités latérales glabres des hanches postérieures largement découvertes: bords latéraux 
rebordés, arqués puis discrètement sinués avant les angles apico-externes; ces derniers 
presque droits, émoussés, peu saillants en arrière, n’atteignant cependant pas le bord 
postérieur du premier segment abdominal: bords postérieurs faiblement concaves et 
pourvus d’une nette encoche proche de la ligne médiane; premier segment abdominal 
pourvu d’une courte carène médiane: le 5ème rebordé; sternites 6 et 7 faiblement 
sclérifiés, rétractés au repos à l’intérieur de l'abdomen. 

Pattes (tarses compris) peu modifiées, les tibias subcylindriques, s’élargissant 
progressivement vers l’apex. Fémur de la patte antérieure (Fig. 5) simple, plus long que 
le tibia, ce dernier plus long que le tarse; protarses sans dimorphisme sexuel: les deux 
éperons apicaux du protibia petits, pratiquement indistincts; mésotibia faiblement 
recourbé, armé sur sa face externe de nombreuses (12-15) épines plus fortement pig- 
mentées; apex avec deux éperons de longueur subégale chez les deux sexes; éperon 
externe fortement modifié chez le mâle (Fig. 8); métatibia coudé à la base, aussi long 
que le tarse, face externe couverte de 12-15 épines effilées et plus fortement pigmen- 
tées, non scindées en deux groupes distincts; apex avec deux éperons dont le plus grand 


LA FAMILLE EUCINETIDAE 129 


atteint la base du peigne apical du segment basal du tarse; ce dernier subégal aux 
articles 2, 3, 4 ou 3, 4 et 5 réunis. 

Edéage (Figs 9, 10) avec le lobe médian tres large, simplement lamelliforme, 
dépourvu de renflure apicale; paramères amplement arrondis apicalement, très 
rapprochés dorsalement. Ce type d’edeage présente quelques analogies avec celui 
d’Eucilodes Vit (Vit, 1977, 1985). 

Biologie inconnue, mais il s’agit d’une espèce récoltée au-dessus de 3000 m. 

Distribution. Connu seulement de la localité-type: Népal, Kandbari district. 

Discussion. Le caractère externe le plus frappant de P. pachys sp. n. est l’aspect 
du tégument: épais, brillant, il est vigoureusement ponctué et exempt de microréti- 
culations et de ponctuation nettement rapeuse. Or la ponctuation franche et grosse n'est 
connue que chez de grandes espèces ailées (Eucinetus Germar, Nycteus Latreille, 
Eucilodes Vit, Euscaphurus Casey), où elle est confinée à la face dorsale de la tête. 
Chez P. pachys elle s’étend, presque identique, sur l’ensemble de la face dorsale et 
ventrale du corps. 

Quant a la conformation de la suture frontoclypéale, formant une sorte de 
gouttiere invaginée, flanquée de chaque coté d’une fossette, elle demeure également 
unique. Sa présence est d’autant plus surprenante qu’on releve, chez les Eucinetidae, 
une tendance certaine a la disparition de la suture frontoclypéale chez toutes les petites 
especes (y compris Bisaya nossidiiformis Reitter) présentant un appareil buccal du type 
modifié. I] pourrait s’agir d’un dispositif rudimenraire du type mycangium (Crowson, 
1981), destiné au transport des spores de champignons (notamment Myxomycetes), 
dont des Eucinetidae sont supposés se nourrir. Cependant des données attestees, 
concernant les particularités alimentaires des Eucinetidae, restent encore trop pauvres 
(Lawrence & Newton, 1980; Chandler, 1991). 


Proeuzkus coecus sp. n. Fig. 2 


Matériel. Holotype 9%, étiqueté: Thailand Doi Inthanon 1750 m. 7.X1.85 Löbl, 
Burckhardt. (MHNG); Paratype. 1 ®, étiqueté: /Thailand-Chiang Mai Doi Saket “1130 m. 4.XII. 
87” P. Schwendinger. (MHNG - exemplaire disséqué conservé en glycérine). 


Wie] 


Espece aptere et anophtalme (femelle), coloration entierement d’un jaune clair 
translucide, épines latérales et cils apicaux des tibias et des tarses fortement rembrunis, 
noiratres; pubescence des téguments fine, couchée. Corps trapu, une fois et demie plus 
longue que large, régulièrement convexe, peu atténué apicalement. Longueur 0,86 mm, 
largeur 0,55 mm, située au niveau posthumeral. 

Téte (Fig. 2). Large (une fois et demie plus large que longue sur la ligne mé- 
diane, labre non compris), modérément convexe, paralléle au niveau des tempes et 
presque semi-circulaire en avant; tempes doublées ventralement, en dessous de l’inser- 
tion antennaire, par une lame latérale dentiforme non prolongée ventralement sous 
forme d’une carène transversale de la tête. Yeux absents; côtés du front fortement 
explanés en avant de l’insertion antennaire; fosses antennaires nulles car non fermées 
ventralement par le bord inférieur explané de la tête, développé chez d’autres espèces. 
Appareil buccal du type piqueur-suceur; suture frontoclypéale absente; clypéus ample- 
ment arrondi, échancré au milieu, finement rebordé; ponctuation forte, éparse, granu- 
leuse. 


130 STANISLAV VIT 


Antennes compactes, deprimees, courtes (plus courtes que la moitié de la largeur 
basale du pronotum), larges (seulement quatre fois plus longues que larges), leur plus 
grande largeur située a peu pres au niveau du Seme segment; articles fermement 
encastres les uns aux autres. Scape visible dorsalement, fortement comprimé, étroit, 
allongé, et protuberant à son bord distal, cachant pariellement le pédicelle; ce dernier 
robuste, subcylindrique,inséré sur la face ventrale du scape; articles 3 a 7 du funicule 
fortement transverses (deux a trois fois plus larges que longs); articles 8 a 10 plus 
allongés, évoquant une massue mal différenciée: segment apical réduit, subovoide, plus 
long que large, surmonte d’une pubescence apicale. 

Base du pronotum doublement sinueuse, fortement projetee en arriere, suplom- 
bant celle des éiytres; scutellum grand mais entièrement caché sous le rebord basal du 
pronotum; côtés droits, finement rebordés; angles postérieurs aïgus, étirés en arrière. 
Ponctuation fine et éparse sur le disque, plus grossière dans les déclivités latérales. 

Elytres d’un tiers plus longs que larges ensemble: côtés arqués, acuminés vers 
l’apex; suture élytrale non soudée; strie juxtasuturale et stries longitudinales absentes; 
six stries vestigiales, rudimentaires et incomplètes demeurent néanmoins perceptibles 
lors d’un examen de l’élytre dans une solution aqueuse. 

Face ventrale avec une forte ponctuation serrée sur les plaques coxales et le 
metasternum; aire médiane des plaques coxales à peine déprimée. 

Epipleures entiers, étroits, atténués à l’apex. Hanches médianes contigües; limite 
médiane du mésosternum obtuse, angle apical renflé, glabre, non étiré en un processus 
médian séparant les hanches: métasternum petit; languette médiane longuement effilée, 
se prolongeant jusqu’au bord postérieur des plaques coxales sous forme d’une fine 
carène médiane: branches latérales étroites, entièrement fusionnées avec les métépis- 
ternes; dispositif de blocage des élytres présent: mésépimères fusionnés avec les 
mésépisternes. Plaques coxales larges mais orientées longitudinalement, laissant décou- 
vertes les extrémités latérales glabres des hanches postérieures: bords latéraux délica- 
tement rebordés, faiblement arqués: angles apico-externes aigus, projetés en arrière, au- 
dessus du bord postérieur du premier segment abdominal; bords postérieurs doublement 
sinueux. Cinq sternites visibles chez la femelle, le cinquième large, ogival, très fine- 
ment rebordé, aussi long que les sternites 3 et 4 réunis. 

Pattes (tarses compris) très robustes, les tibias déprimés, s’élargissant fortement 
vers l’apex. Protibia aussi long que le tarse, apex sans éperons apicaux distincts; fémurs 
des pattes antérieures et médianes avec une saillie dentiforme contigüe au trochanter; 
face externe du mésotibia avec deux rangées longitudinales, serrées, de soies spini- 
formes plus pigmentées: apex avec deux éperons effiles; métatibia aussi long que le 
tarse, face externe du metatibia garni de nombreuses soies spiniformes rembrunies, for- 
mant un groupe préapical plus ou moins nettement séparé, apex avec deux éperons 
effilés plus longs que le segment basal du tarse; ce dernier plus court que les articles 2, 
3, 4 ou 3, 4 et 5 réunis. 

Distribution. Les deux captures proviennent de la province de Chiang Mai au 
nord de la Thailande, de tamisages de débris végétaux en forét de montagne, entre 1200 
et 1700 m. 


LA FAMILLE EUCINETIDAE 131 


Fics 6-11 


Proeuzkus pachys sp. n. : 6, appareil buccal, vue ventrale; 7, antenne; 8 mésotibia et les éperons 
apicaux (d); 9, édéage, vue ventrale; 10, édéage, vue latérale; 11, segments abdominaux VI et 
VII (3). (Mesures données en mm). 


132 STANISLAV VIT 


Discussion. L’ anophtalmie de cette espèce et la présence d’un puissant appareil 
buccal du type piqueur-suceur ont pour corollaire de tres notables modifications dans la 
morphologie de la tête: de longues tempes parallèles: bords latéraux du front très 
fortement explanés; échancrure gulaire occupant pratiquement toute la face ventrale de 
la tête, d’où la suppression du pli transverse de la tête, dont il ne subsiste que les 
extrémités latérales dentiformes: absence du bord latéral inférieur de la tête qui 
normalement constitue la paroi ventrale de la fosse antennaire. D’autres modifications à 
caractère convergent, que l’on retrouve chez d’autres espèces et genres de la famille: 
fusion des pièces sternales, absence de la strie juxtasuturale, réduction des stries 
élytrales ou réduction du nombre de sternites apparents, sont à mettre en corrélation 
avec la réduction notable du corps (0,86 mm = la plus petite espèce connue). Il serait 
utile de pouvoir connaître le mâle et la structure de l’édéage de cette espèce. 


Tohlezkus laticanthus sp. n. Figs 12-16 


Matériel. Holotype d , étiqueté: MALAYSIA: Pahang, 4 mi NE Cameron Highlands, 23- 
25.IV. 1977, (berlese leaf litter), L. Watrous. (CUIC). 

Espece aptere (pas d’ailes distinctes par transparence des téguments chez le 
male), coloration entièrement d’un brun roussatre clair, côtés de la tête rembrunnis, cils 
apicaux des tibias et des tarses a peine plus foncés: pubescence fine, couchée. Corps 
allongé, deux fois plus long que large, faiblement acuminé apicalement. Longueur 1,05 
mm, largeur 0,62 mm, située au niveau posthumeral. 

Tete. Large, subogivale, modérement convexe, parallele au niveau des tempes; 
côtés convergeant en avant; clypéus arrondi; ponctuation forte, subräpeuse. Yeux bien 
développés, latéraux, formés d’un gros grain hyalin fortement saillant latéralement; 
limites latérales du cranium fortement pigmentées; insertion antennaire entierement 
cachée sous le bord explané du front; cavité antennaire fermée ventralement. Appareil 
buccal du type piqueur-suceur; suture frontoclypéale absente; bord antérieur reborde. 

Antennes compactes, peu déprimées, un peu plus longues que la moitié de la 
largeur basale du pronotum, épaisses, au moins 5 fois plus longues que larges, leur plus 
grande largeur située au niveau du 8ème segment; articles encastrés les uns dans les 
autres. Scape en partie caché sous le rebord explané du front, cylindrique; pédicelle 
cylindrique, subcarré, inséré axiallement; articles 3 à 6 du funicule courts, transverses 
(deux fois plus larges que longs); articles 7 a 11 plus robustes, plus allongés, faiblement 
transverses; segment apical grand, subovalaire, plus long que large, surmonté de 
pubescence apicale. 

Base du pronotum doublement sinueuse, projetée en arrière, suplombant la base 
des élytres; scutellum petit, entièrement caché: côtés très finement rebordes; angles 
postérieurs aigus, étirés en arrière. Ponctuation nette sur le disque, plus grossière dans 
les déclivités latérales. 

Elytres une fois et demie plus longs que larges ensemble, ratio Lo.s./La. = 1,46: 
côtés peu arques, subparallèles, faiblement acuminés; vraisemblablement soudés sur la 
suture; strie juxtasuturale oblitérée; stries longitudinales absentes: cinq stries ponctuées 
vestigiales et incomplètes cependant perceptibles dans la portion basale lors d’un 


LA FAMILLE EUCINETIDAE 


Fics 12-15 


Tohlezkus laticanthus sp. n.: 12, édéage, vue ventrale; 13, segments abdominaux VI et VII (£): 


14, hanche antérieure et profémur (normalement constitués): 15, patte antérieure gauche (térato- 
logique). Mesures données en mm. 


134 STANISLAV VIT 


examen dans une solution aqueuse. Ponctuation tres serrée et subräpeuse, arrangee, pres 
de la base, partiellement, en paliers transverses . 

Face ventrale entierement, densement, et räpeusement ponctuée; aire médiane 
des plaques coxales nettement déprimée. 

Epipleures entiers mais de forme particulière: d’abord étroits et fortement 
comprimés dans leur portion post-humérale, puis larges, s’élargissant même progressi- 
vement jusqu'au niveau du troisième sternite et brièvement atténués vers les angles 
suturaux. Processus mésosternal semblable à celui rencontré par exemple chez Eusca- 
phurus spinipes Vit; élancé, cordiforme, pourvu d’une carène médiane, acuminé apica- 
lement, mais ne séparant pas les hanches médianes; cavités cotyloïdes contigués. 
Métasternum petit; languette médiane fortement éffilée, se prolongeant jusqu’au bord 
postérieur des plaques coxales sous forme d’une fine carène médiane; branches latérales 
étroites, tout au plus faiblement élargies à leurs extrémités latérales; entièrement 
fusionnés avec les métépisternes; mésépimères, fortement transverses, subfusionnés, 
leur suture avec le mesépisterne distincte: mésosternum et mésépisternes fusionnés. 
Plaques coxales orientées longitudinalement, étroites, laissant largement découvertes 
les extrémités latérales glabres des hanches postérieures: bords latéraux délicatement 
rebordés, faiblement arqués; angles apico-externes aigus, émoussés au sommet, forte- 
ment projetés en arrière au-delà du bord postérieur du premier sternite; bords posté- 
rieurs fortement convergents vers la ligne médiane. Cinq sternites visibles ( sûrement 
chez les deux sexes), le cinquième large, ogival, finement rebordé, aussi long que les 
sternites 3 et 4 réunis. Chez le mâle les sternites 6 et 7 (Fig. 13) restent faiblement 
sclérifiés et entièrement retractés dans l'abdomen. 

Pattes (tarses compris) robustes, déprimées, tibias s’elargissant fortement vers 
l’apex; profémur (Fig. 14) avec une saillie dentiforme contigüe au trochanter et un 
peigne de soies longeant le bord externe: patte tératologique (cf. Remarque) de cette 
exemplaire (Fig. 15) présente un éperon apical et le protarse non dilatés chez le mâle 
(!); mésotibia faiblement incurvé, plus court que le tarse; face externe avec deux ran- 
gées longitudinales serrées de soies spiniformes, rembrunies, apex avec deux éperons 
dont l’interne fortement développé chez le mâle, épais, incurvé et renflé apicalement, 
presque aussi long que le segment basal du tarse: métatibia faiblement incurvé, aussi 
long que le tarse; face externe garnie de nombreuses soies spiniformes plus pigmentées 
et d’un peigne préapical d’épines foncées, apex avec deux éperons dont un remar- 
quablement développé, en forme de “chausse-pied” (Fig. 16), dilaté, spatuliforme, 
tronqué apicalement, plus long que le segment basal du tarse; ce dernier subégal aux 
segments 2, 3, 4 ou 3, 4 et 5 réunis. 

Edeage (Fig.12) pourvu de structures membraneuses internes et de parameres 
explanés du côté dorsal et finement ciliés a l’apex. 

Caracteres sexuels. Un seul male étant connu, il n’est pas possible de définir si 
l’éperon apical modifié des métatibia (Fig.16) est, ou n’est pas, un caractère sexuel du 
mâle (au même titre que celui des mesotibias), ou un caractère propre de cette espèce. 

Remarque. Les pattes antérieures de cet exemplaire unique sont incomplètes 
(Figs 14-15). A l’une manque le tibia, l’autre est sujette à une malformation curieuse, 
où son tibia, de surcroît malformé, est articulé directement avec la hanche. 


LA FAMILLE EUCINETIDAE 135 


Discussion. Les deux caractères remarquables de cette espèce sont ses épi- 
pleures dilatés dans la partie subapicale et I’hypertrophie de l’éperon apical des méta- 
tibias. On relève néanmoins, ou niveau de la famille, plutôt une tendance à la réduction 
des épipleurs et une régression des éperons apicaux des tibias. 


Eucinetus xaca Sp. n. Figs 17-20 

Matériel. Holotype d , étiqueté: Mexico: Oaxaca, 12 km N Oaxaca City, hwy 175 1900 
m. 28.IX. 1990, sifting litter at small stream, tropical mountane forest. (ZML). Paratypes. 22 9 
mêmes données que l’holotype; 1d (1.X. 1990), 29 © (5.X. et 8.X.1990) Mexico: Oaxaca, 12 
km N Oaxaca City, hwy 175 1900 m. 1.-8.X.1990 leg. R. Raranowski, Pit-fall traps near small 
stream, trop. mount. forest. (ZML); 25 4, 29 2 Mexico: Oaxaca 12 km N Oaxaca 2000 m., 
15.IX.1986 R. Baranowsk, sifting litter in creek, trop. mont. forest; 26 4,29 © Mexico: Oaxaca 
10 km. N Oaxaca 1900 m. 13.1X. 1986 R. Baranowski, sifting litter at small stream, trop.mont. 
forest. (ZML, CSV). Autres. 1% (xaca ?) Jalisco 50 km S Puerto Vallarta 800 m. 16.V1.1986 leg. 
M. Sörensson & B. Martensson (ZML). 

Longueur dorsale médiane (tête non comprise) 3,1-3,6 mm, corps subparallele, 
la plus grande largeur 1,73-1,86 mm; ratio Lo.m. (sans téte)/ La. = 1,97-2,01 Coloration 
presque uniforme, d’un brun rougeätre foncé, brillant, partie apicale des élytres parfois 
sombre: pattes plus claires: antennes tricolores, claires et noirätres. 

Tête. Transverse, longueur médiane inférieure à la largeur du front entre les 
yeux, ratio Lo.m./La.F. = 0,83-0,92; ponctuation fine, serrée. 

Antennes tricolores (articles 1-3 bruns, 4-9 noirätres, 10 et Il jaunätres), 
longues environ comme la base du pronotum mesurée en vue dorsale, ratio Lo.A./La.P. 
= 0,95-1,11; tous les articles allongés; pédicelle trois fois plus long que large, plus long 
que le 3ème; ce dernier grêle, tout juste plus court que le 4ème; articles 4 à 10 
subégaux, diminuant progressivement en longueur; | 1ème article une fois et demie plus 
long que large. 

Ponctuation du pronotum fine et serrée sur le disque, aussi forte que celle de la 
tête dans les déclivités latérales. 

Elytres longuement subparallèles, ratio Lo.m./La.(sexes confondus) = 1,1,60- 
1,74, rebord latéral perceptible de dessus au moins dans le tiers apical. Strie juxta- 
suturale oblitérée a environ trois longueurs de scutellum de la base; interstrie juxta- 
suturale faiblement enflé à l’apex; stries longitudinales seulement peu visibles, mar- 
quées tout au plus dans la moitié apicale, interstries à peine convexes; ponctuation fine, 
paliers transverses très serrés (4 paliers = 0,1 mm). 

Face ventrale. Epipleures raccourcis, se confondant avec le bord de l’élytre au 
niveau du 4e sternite; leur largeur après l’étranglement posthuméral égale à la moitié de 
la longueur du 3ème article antennaire. Metasternum densement ponctué; languette 
métasternale longue, subtriangulaire, suture médiane nettement imprimée; sutures trans- 
verses distinctes. Mesurés sur leur bords externes, les métépisternes sont aussi longs 
que les mésépimères, ces derniers presque aussi larges que longs; suture séparant les 
mésépimères des mésépisternes nette, non renflée. Plaques coxales larges, transversales, 
cachant parfaitement les extrémités latérales des hanches postérieures; côtés non 
rebordés: angles apico-externes arrondis (atteignant le niveau du bord postérieur du 
premier segment abdominal); bords postérieurs simples, convergeant vers la ligne 
médiane. Bord postérieur du Se et 6e sternite amplement échancré (à ). 


186 STANISLAV VIT 


TI RER 
RR a 


TTS! 


/ 


SS 


pri 


PAZZO 


urn! Zu 
Stino 


Fics 16-20 


Tohlezkus laticanthus sp. n.: 16, éperon apical des métatibias; Eucinetus xaca sp. n.; 17, édéage, 
vue ventrale; 18, édéage, détail de la partie apicale du lobe médian: 19, édéage, vue latéral du 
méme; 20, patte postérieure, tibia et tarse. Mesures données en mm. 


LA FAMILLE EUCINETIDAE 137 


Patte antérieure.Tarse fortement dilaté chez le male; subégal chez les deux sexes 
à la longueur du tibia, ce dernier modérément renflé chez le mâle, subparallele chez la 
femelle. 

Patte médiane. Tibia très nettement plus court que le tarse, progressivement 
élargi, bord externe avec 3 à 8 épines noires, couchées, apex avec un seul éperon peu 
développé chez la femelle, deux éperons de longueur subégale chez le mâle, l’éperon 
interne simplement épaissi. 

Patte postérieure (Fig. 20). Tibia nettement plus court que le tarse, légèrement 
incurvé, garni sur sa face externe de 13 - 20 épines semi-dressées, plus 2 - 4 épines 
dans le groupe pré-apical pas nettement singularisé; un seul éperon apical, aussi long 
que la moitié du basitatarse; longueur de ce dernier nettement inférieure à celle des 
articles 2, 3, 4 ou 3, 4 et 5 réunis. 

Edéage (Figs 17-19) pourvu de puissantes excroissances dentiformes à la base 
de l’étranglement subapical du lobe médian, portion distale des paramères lamelliforme 
(cf. Discussion). 

Distribution. Mexique méridional, Oaxaca. 

Discussion. Avec d'importantes modifications de la portion apicale des para- 
mères (normalement arrondie) et de la portion subapicale du lobe médian (normalement 
tubuliforme), muni ici de puissantes excroissances latérales fortement sclérifiées, 
l’édéage de Eucinetus xaca sp. n. confirme une évolution morphologique tout a fait 
interessante. Ce type d’édéage a déjà été signalé (Vit, 1990) pour deux autres espèces 
du groupe haemorrhoidalis, à savoir: E. pecki Vit et E. apterus Vit, décrites également 
du Mexique. Or cette évolution vers une forme différente de l’édéage - pour l'instant 
parfaitement isolée au sein du genre Eucinetus - apparaît chez des espèces mexicaines 
de façon tout à fait endémique. Elle ne se rencontre ni au nord, chez les espèces néarc- 
tiques, ni au sud, chez celles néotropicales. L’édéage d’E. xaca Sp. n. est très proche du 
celui d’E. apterus, l'unique espèce aptere du groupe haemorrhoidalis, bien caractérisée 
par ailleurs par une réduction des métépisternes, relative à la disparition des ailes 
métathoraciques. 


REMERCIEMENTS 


Je tiens à remercier ici Dr C. Besuchet et Dr I. Löbl du Muséum d'Histoire 
naturelle de Genève, Dr A. Smetana d'Ottawa, Dr Quentin D. Wheeler du Département 
d’entomologie de l’Université Cornell d’Ithaca et Dr R. Danielson du Muséum de 
Zoologie de Lund, pour m'avoir confié l’étude des matériaux traités dans cette note. 


BIBLIOGRAPHIE 


CROWSON, R. A. 1981. The Biology of the Coleoptera. Academic Press, London, 802 pp. 


CHANDLER, D. S. 1991. Comparison of some slime-mold and fungus feeding beetles (Coleoptera: 
Eucinetoidea, Cucujidoidea) in an old-growth and 40-year-old forest in New Hampshire. 
The Coleopterists Bulletin 45(3): 239-256. 


LAWRENCE J. F. & NEWTON A. F. Jr. 1980. Coleoptera associated with the fruiting bodies of 
slime molds (Myxomycetes). The Coleopterists Bulletin 34(2): 129-143. 


138 STANISLAV VIT 


Vit, S. 1977. Contribution à la connaissance des Eucinetidae (Coleoptera). Revue suisse de 
Zoologie 84: 917-935. 

VIT, S. 1981. Une nouvelle espèce du genre Tohlezkus Vit de Taiwan (Coleoptera, Eucinetidae). 
Revue suisse de Zoologie 88: 769-774. 

Vir, S. 1985. Etude de la morphologie des espèces paléarctiques du genre Eucinetus Germar et 
quelques remarques sur son utilisation taxonomique (Coleoptera: Eucinetidae). Revue 
suisse de Zoologie 92: 421-460. 

Vit, S. 1990. Revision des espèces néotropicales du genre Eucinetus Germar (Coleoptera: Euci- 
netidae). Naturaliste canadien (Revue d’Ecologie et de Systématique) 117: 103-122. 

Vit, S. 1995. Deux especes nouvelles d’Eucinetidae d’ Amérique du Nord particulierement inte- 
ressantes (Coleoptera: Eucinetidae). Elytron 9: 125-137. 


REVUE SUISSE DE ZOOLOGIE 107 (1): 139-151; mars 2000 


Spatio-temporal distribution of size classes and larval instars of 
aquatic insects (Ephemeroptera, Trichoptera and Lepidoptera) in a 
Potamogeton pectinatus L. bed (Lake Geneva, Switzerland) 


Ruth BÄNZIGER 
Laboratoire d'écologie et de biologie aquatique, 18 ch. des Clochettes, 
University of Geneva, CH-1206 Geneva, Switzerland. 


Spatio-temporal distribution of size classes and larval instars of aquatic 
insects in a Potamogeton pectinatus L. bed (Lake Geneva, Switzerland). - 
Temporal changes of aquatic insect instars or size classes were monitored ın 
different parts of a Potamogeton pectinatus bed. The hypothesis of a diffe- 
rent distribution of the aquatic insects in the macrophyte bed according to 
their life stage and of a spatio-temporal segregation of congeneric species 
was tested. Head capsules widths of seven insect species (Caenis horaria 
and C. luctuosa (Ephemeroptera), Mystacides azurea, M. longicornis, 
Oecetis lacustris and O. ochracea (Trichoptera) and Acentria ephemerella 
(Lepidoptera)) were measured at monthly intervals from May to November 
1994. Samples were taken in the edge and in the centre of the macrophyte 
bed on all sampling occasions, and in May, June and July, additional samples 
were taken from the 2 m sediment belt adjacent to the macrophyte bed. Each 
couple of congeneric species showed segregation by size before hibernation 
and showed delayed emergence patterns. 


Key-words: Lake - Ephemeroptera - Trichoptera - Lepidoptera - macro- 
phyte - distribution - size class. 


INTRODUCTION 


Habitat segregation or space partitioning among closely related species have 
been often addressed in rivers (Malas & Wallace, 1977; McAuliffe, 1984). Few papers, 
however, have been concerned with this subject in lakes. Species with similar 
ecological niches are not always separated in space and time, thus competition is not 
necessarily involved and some congeneric species do coexist (Hildrew & Edington, 
1979). Segregation, if there is any, is therefore likely to occur at another scale than at 
the species level. In larval insects or nymphs, delayed growth of the last instars or 
nymphal stages have been observed, often resulting in temporal segregation of 
emergence (Macan, 1965; Tudorancea & Green, 1975; Malas & Wallace, 1977; 
Bengtsson, 1981; Sweeney & Vannote, 1981; Brittain, 1982). Hildrew & Edington 
(1979) showed that two congeneric hydropsychid caddisflies avoided coexistence by 
different microhabitat colonization of some instars. Hydropsychidae were also studied 


Manuscript accepted 13.09.1999 


140 RUTH BÄNZIGER 


by Muotka (1990) who showed them having different microhabitat preferences accor- 
ding to different larval stages. Minshall (1984) observed changes in spatial distribution 
of insects according to developmental stages. Altogether, between egg and adult, the 
growing insects may live through many different feeding modes, behaviour or habitat 
niches (Winterbourn, 1971; Resh, 1979; Palmer et al., 1993). These changes according 
to development allow them to reduce competition if resources are scarce. 

In a previous paper (Bänziger er al., subm.), we demonstrated differences in 
densities of several invertebrate taxa between the edge and the centre of macrophyte 
beds. The question arose whether these differences in density could be related to larval 
instar or size class distribution. 

The purpose of the present study was to test the following hypotheses in a 
Potamogeton pectinatus L. bed of the littoral zone of Lake Geneva: 1) distribution of 
insects in macrophyte beds changes according to larval instar or size class; and ii) 
congeneric species living in the same macrophyte bed differ in size and/or timing of 
larval instars. 


MATERIAL AND METHODS 


The samples were taken in a Potamogeton pectinatus L. bed near Corsier 
(46°16' N, 6°12' E) in the littoral zone of Lake Geneva, Switzerland. 

Samples were taken by scuba diving at a depth of 3.0 - 3.5 m in the edge, in the 
centre and adjacent to the macrophyte bed. The edge was defined as the margin of the 
bed characterised by lower macrophyte stem density, the presence of more filamentous 
algae (depending on the season) and by shorter shoots than in the centre of the bed. 
This corresponded to the 2 m wide outer belt of the macrophyte bed. Macrophyte 
density in the centre of the bed was 80 stems m’? at maximum density, i.e. in July. The 
(arbitrarily) 2 m wide area around the macrophyte bed, consisting of sediment, was 
defined as the adjacent sediment. 

Macrophyte samples were taken from May through to November 1994, adjacent 
sediment was sampled from May to July. When the macrophytes senesced - from 
October to November - sampling was carried on in the plant underlying sediments and 
the remains of the Potamogeton bed. Collections were made using different sampling 
gears depending on the substrate: 

Collections of fully grown macrophytes were made using a sampler modified 
after Gerking (1957) with a surface area of 0.25 m? and a height of 1 m. When 
vegetation was less dense a frame of 0.0625 m? with an attached net was used. Each of 
the two sampling gears was lifted down on the macrophytes by a scuba diver, the plants 
were teamed out of the sediment and the sampler was closed: the Gerking-like sampler 
by a trap and the net by a string. 

Sediments underlying the macrophyte bed were collected using corers covering 
an area of 0.005 m? and pushed 10 cm deep into the sediments. Adjacent sediments 
were collected inside a 0.25 m? frame using the same net as for vegetation samples. 

In the laboratory, samples were thoroughly rinsed with tap water and the 
macroinvertebrates were retained in a 250 um sieve. They were conserved in 4% 
formalin. 


AQUATIC INSECTS IN MACROPHYTE BEDS 141 


Seven insect species were retained for measurements: Caenis horaria (L.), C. 
luctuosa (Burm.) (Ephemeroptera), Mystacides azurea (L.), M. longicornis (L.), Oecetis 
lacustris (Pictet), O. ochracea (Curtis) (Trichoptera) and Acentria ephemerella (Denis 
and Schiffermiill.) (Lepidoptera) on the basis of their abundance in the samples, their 
identification and/or the availability of congeneric species. 

Larvae of mayflies were assigned to size classes as they do not have easily 
distinguishable cohorts (Benke & Jacobi, 1986). The size classes of Caenis spp. and the 
instars of Trichoptera and Lepidoptera were assigned on the basis of head capsule 
widths (HCW) as they were often reported to be more reliable than body length in 
separating the different stages (Bradbeer & Savage, 1980; Bass ef al., 1982). 

Measures of HCWs were made at their widest point including the eyes (Smock, 
1980; Bass er al., 1982). All measures were made with an accuracy of 0.025 mm using 
a dissecting microscope with a micrometer. 

Head capsules widths of Caenis spp. were divided into 18 classes of 0.05 mm 
each (Table 1). Larvae of Mystacides spp. and Oecetis spp. were divided into five larval 
instars (Table 2). At instar I, genera could not be keyed to species, so they were termed 
“juveniles”. 


TABLE | 
Correspondance between size classes and head capsule widhts (HCW) of Caenis spp. 


size classes HCW of Caenis size classes HCW of Caenis size classes HCW of Caenis 


spp. (mm) spp. (mm) spp. (mm) 
I 0.125 - 0.2 U 0.575 - 0.65 13 1.025 - 1.1 
2 0.2 = 0.275 8 0.65 - 0.725 14 ILES 
3 0.275:- 0.35 9 0.725 - 0.8 15 12.1779%-.1:25 
+ 0.35 - 0.425 10 0.8 - 0.875 16 E25 les 25) 
5 0.425 - 0.5 11 0.875 - 0.95 17 1.325 - 1.4 
6 12 


050575 2 0:9377.1:025 18 1.4 - 1.45 


Data in the literature dealing with HCWs of A. ephemerella were scarce. Five 
instars were reported in the literature and Haenni (1974, 1980) identified the sizes of 
the different instars by collecting individuals in the field and by rearing them. 


TABLE 2 


Correspondance between instars and head capsule widhts of Mystacides azurea, M. longicornis, 
Oecetis lacustris and O. ochracea. 


instars HCW of instars HCWofM. instars HCW of instars HCW of O. 
Mystacides longicornis Oecetis lacus- ochracea 
azurea (mm) (mm) tris (mm) (mm) 


fovea O25 0175" uv  O125 DIS uva 7025-015 juve) :09925520:2 

II 0.175- 0.2 I 0.175 - 0.2 Il 0.25 - 0.3 
IT 02-0275 I 0252025 II 0.25 - 0.35 II 0.375 - 0.475 
MV 0325-04 IV 0.375-0.475 IV 0.4 - 0.525 IV 0.625 - 0.825 
V 0.5 - 0.675 V 0.575 - 0.75 V 0.675 -0.875 V OOS) Sal A275) 


142 


= 10% 


Relative abundance of size classes (%) 


RUTH BANZIGER 


31.V.94 Caenis horaria, n = 68 
C. luctuosa, n = 45 


nine 


04.V11.94 
Caenis horaria, n = 3 


C. luctuosa, n = 56 | | E 


05.V111.94 Caenis horaria, n = 87 
| C. luctuosa, n = 116 
4 | IT Pes 
26.V111.94 


Caenis horaria, n = 335 
C. luctuosa, n = 334 


19.1X.94 


TIR 
31.X.94 
i Pdl | | | Île 
29.X1.94 
alli nino 
8 ‘OR Ne 


2 = 6 


Caenis horaria, n = 435 
C. luctuosa, n = 693 


Caenis horaria, n = 319 
C. luctuosa, n = 203 


Caenis horaria, n = 279 
C. luctuosa, n = 207 


2 14 16 18 Size classes 


AQUATIC INSECTS IN MACROPHYTE BEDS 143 


Combining his results and our measurement yielded the following correspondence 
between HCWs and instars. Instar I: 0.2-0.3 mm; H: 0.325-0.5 mm; III: 0.525-0.775; 
IV: 0.8-0.925; V: 0.95-1.1 mm. The different instars are overlapping. 

Emergence time of larvae and nymphs was based on numbers and size of the 
larvae and nymphs at the different sampling sessions. 


RESULTS 


COMPARISON OF GROWTH AND EMERGENCE TIME OF CONGENERIC SPECIES 


The three pairs of congeneric species Caenis spp., Mystacides spp. and Oecetis 
spp. showed staggering in emergence patterns. This delay was already prepared for in 
autumn, C. luctuosa, M. longicornis and O. lacustris did stop growing from the middle 
of September, whereas, C. horaria, M.azurea and O. ochracea showed some growth 
until October - November. 

According to the low numbers catched beginning of July, imagos of Caenis 
horaria emerged between end of May and June. Juveniles appeared in the samples at 
the beginning of August, but their abundance was highest at the end of August (Fig. 1). 
At that time, the new generation extended over one month divided into two density 
peaks: the first one (low density) ranging from size classes 9 to 15 and the second one 
(high density) ranging from 2 to 8. The nymphs from the first (9-15) peak emerged by 
the middle of September. Nymphs of the second peak grew until the end of October. 
Most of them entered the winter period at size classes from 8 to 11. 

Caenis luctuosa emerged later than C. horaria: at the beginning of July. 
Juveniles were collected in the samples at the beginning of August, as for C. horaria. 
The two species followed the same growth schedule (with two size class peaks at the 
end of August) until the middle of September. At that time C. luctuosa stopped 
growing, thus entering the winter period at size classes from 3-8 essentially. 

Interpretation of the growth of Mystacides azurea was more difficult as 
sampling in May seemed to have bypassed the emergence of M. azurea (Fig. 2). M. lon- 
gicornis emerged at the beginning of July and the first hatched larvae of M. azurea 
were ready to emerge at the beginning of August. Some M. longicornis larvae did also 
reach instar V at the end of August and emerged or disappeared until the middle of 
September. By the end of October, instars II to IV of M. longicornis and III and IV of 
M. azurea were present. M. longicornis did not grow further, whereas M. azurea was 
found from instars II to V at the end of November. Entering the winter in the last instar 
could preclude to an early emergence in spring. 

In Oecetis spp., emergence time extended from the end of May through August 
for O. ochracea, while the emergence of O. lacustris was observed from beginning of 
July through August. Some of the first hatched larvae of the two species appeared at the 


Fic. | 


Size classes and headcapsule widths of Caenis horaria (open columns) and C. luctuosa (dark 
columns). Black arrows: emergence. 


144 


RUTH BÄNZIGER 


beginning of August. O. lacustris did not grow further from the middle of September 
and overwintered in the instars II and III, while growth of O. ochracea lasted until the 
end of October and it overwintered in the instars III and IV. 


GROWTH AND EMERGENCE OF ACENTRIA EPHEMERELLA 


End of May was the end of the emergence period for A. ephemerella (Fig. 2). 
There were still some pupae found. At the beginning of July, the first new larvae 


Instars ( 


Bu Bi wn NN 


Mystacides azurea 


Mystacides longicornis 


= a cone a à 


Ov) 


RE - 180 # 


ne=e23 n= 19 


i= SSD Lr cia gaze ag 


a 31.V.94 04.V11.94 O5.VIII.94 26.V111.94 19.1X.94 31.X.94 29.X1.94 
° 
SS 7 

2 Instars (Mouv Ki I NV DV) 
7) 
= 
A, Oecetis ochracea 
oO 
n= 10 fae if n= 116 # = 128 1 
a | 
le: = Lal 
° 
o 
© 
c 
3 
E Oecetis lacustris 
ol) Wee el ee end 
o 
2 
Kc: 
© 
cc 31.V.94 04.V11.94 O5.VIII.94 26.VIII.94 19.1X.94 31.X.94 29.X1.94 

Instars ( J I Zu Ww By) 

— ee Br =a 

mmm 25% | 
| Acentria ephemerella 
n= 109 n= 191 n= 268 Dell n = 23 


I A 
= 


31.V.94 05.V111.94 


04.V11.94 


26.VIII.94 19.K.94 31.X94 29.X1.94 


Fic. 2. Frequency of the instars of five insect species (Mystacides azurea, M. longicornis, 


Oecetis lacustris, O. ochracea and Acentria ephemerella). 


Juv.: indetermined juveniles of 


either Mystacides spp. or Oecetis spp. Black arrows: emergence. 


AQUATIC INSECTS IN MACROPHYTE BEDS 145 


appeared and at the beginning of August, all larval instars were present. The question 
remained whether instar V larvae were the first hatched of the 1994 cohort or the last 
hatched of the 1993 cohort. From August to November, young larvae grew to instar II 
and III and instar IV larvae disappeared. Abundance of instar II and IH larvae was 
similar at the end of November. 


SPATIAL DISTRIBUTION OF INSTARS 


None of the seven insect species studied showed significant differences in 
relative abundance of the different instars or size classes between edge and centre of the 
macrophyte bed and the adjacent substrate. Our data and figures did however show 
higher relative abundances of instar V larvae on the plants (edge and centre), especially 
for Oecetis spp. and Mystacides spp. (Fig. 4). 

Caenis spp. showed no tendency to colonize either plants or adjacent sediments 
(Fig. 3) and Acentria ephemerella was found quite exclusively on the plants (Fig. 5). 
M. azurea was slightly more abundant on the adjacent sediments and on the plants in 
the edge at the beginning of August, whereas M. longicornis was more abundant in the 
centre at instar V. Thus, it seemed to emerge in higher densities from the centre of the 
macrophyte bed than M. azurea. 

Oecetis spp. were the only species which were quite abundant in the sediments. 
In May, they were mostly emerging from the sediments and in the subsequent sampling 
periods they were present in higher densities on the macrophytes, mainly at instar V. 
The bulk of emergence of O. lacustris originated from the edge, while O. ochracea 
emerged indifferently from the edge or the centre of the macrophyte bed. 

Acentria ephemerella showed highest densities throughout the sampling sessions 
in the centre of the macrophyte bed, but at the end of May it was mainly found in the 
edge, besides of the pupae which were found in the centre. Only one individual was 
found at the beginning of August on the adjacent sediments. 


DISCUSSION 


The distribution of larval instars and size classes of the investigated congeneric 
insect species showed that temporal spacing of emergence was a major factor involved 
in their segregation. Indeed, several authors pointed out the importance of separated 
swarming in order to optimise mating success (Brittain, 1982). Moreover, the instar 
segregation already took place in autumn which allows to minimise resource depletion 
by similarly sized and similarly feeding larvae in winter when food sources are scarce. 
In Oecetis spp., this difference was enhanced by the lower size of O. lacustris (usually 
instar V larvae of O. lacustris had the size of instar IV larvae of O. ochracea). Size 
differences between species were lower for the two other congeneric species studied 
(Mystacides spp. and Caenis spp.). 

The spacing of emergence involved that some species (1.e. Oecetis spp.) emer- 
ged before macrophyte resumed growth and they therefore did not need the presence of 
plants to complete their development. However, once the macrophytes were well 
established, Oecetis spp. and Mystacides spp. were found on macrophytes prior to 


RUTH BÄNZIGER 


146 


‘JUDLUIPas 
juaovlpe ai uo pue poq a}Aydosov ou} JO 2HU99 OY) ur pur 28P9 AU} ur PSOMON] 7) PUB VLUYLOY SIUAD) JO SISSLII 9ZIS JUIIOJJIP JO Aguonbo.y 


€ Old 


sessejd aZIS8LZ/LILSLYLELZLILOL Bee ee! ERZIELEN. 
~ BE 


4 
4 
4 
4 
4 
4 
4 
4 
4 
4 
# 
4 
4 
CÀ 
4 
4 
4 


ve IIIA SO r6 'IIlA SO 


Ol LVOU SN CUT ei EOIN Gath ye. ee tzid BELZELOLStPLetct tt Ot 
A 4 a 


IAETOTTE- 
ESS 


r6lIAPrO 


VE lA vO 


LABO EGIZIA ERO 8698 ee: RATTO [Re ez IRL CANON GO SOC ze I 
7 "Eee aeg DT 
À H 


(%) Sessejo azis jo sauepunge anıjejay 


paq }uejd jo 811U99 | | Gz 


pag }uejd jo abpa ZA 
Juswıpas Juaoelpe EJ 


n 
qd 
+ 
4 
’ 
[1 
è 
à 
A 
4 
3 
# 
# 
CA 
4 
[A 


089 


esonjon| SIU9E) euesoy sijuaeg 


AQUATIC INSECTS IN MACROPHYTE BEDS 147 


emergence. This may be ought to the advantage of the closer distance to the air-water 
interface which makes emergence easier and reduces the danger of predation (Rooke, 
1984). 

Among the seven species studied only two are known to mostly rely on macro- 
phytes for feeding: Mystacides longicornis and Acentria ephemerella. Thus the main 
food resource in relation with macrophytes (1.e. plant tissue and periphyton) was not 
competed for to complete development (Berg, 1941; McGaha, 1952; Lepneva, 1966). 
The absence of difference in spatial distribution of congeneric taxa may either indicate 
that there is no spatial segregation or that it occurs at the microhabitat scale, as 
observed by Hildrew & Edington (1979) in rivers, rather than at the edge and centre 
scale of macrophyte beds. As Magdych (1979) and Miiller-Liebenau (1956) reported, 
spatial segregation may take place between leaves, stems and roots at different heights 
(top, middle, bottom) of the plant. 

Some instar I larvae of Oecetis spp. and Mystacides spp. may have been over- 
looked, thus biasing abundance data on the distribution of juveniles (Bass et al., 1982). 

Caenis spp. showed higher densities on the sediments. They emerged from any 
substrate and except for emergence and hibernation, their size classes were synchro- 
nised. The separation of the newly hatched nymphs into a fast growing (emerging after 
two months) and slow growing (hibernating) generation was already evidenced for 
C. horaria by Oertli (1992) in ponds of the Geneva area. Landa (1968) observed the 
same pattern for C. horaria during its study on central Europe Ephemeroptera. The 
variable life cycle patterns of Caenidae were reported by many authors (see Clifford, 
1982). Thus, it is interesting to note at least identical patterns in different years and 
biotopes, but in the same area (i.e. Geneva), for one species. 

Oecetis spp. illustrated the movement of larvae between the sediments and the 
macrophytes. Last instar larvae were more abundant on the sediments in May than in 
the subsequent sampling periods. It is likely that the larvae of O. ochracea did not have 
enough time (or food) to colonize the macrophytes early in the season and therefore 
they were still found on the adjacent sediments at the time of emergence. O. lacustris 
which emerged at the beginning of July, was able to emerge partly from the macro- 
phytes. O. ochracea seemed to be clearly bivoltine, whereas O. lacustris showed a 
mainly univoltine pattern. These two species are at least partly predators so they are not 
relying on macrophytes for feeding (Mackay & Wiggins, 1970). 

M. longicornis, which was found in substantially higher densities on macro- 
phytes than on sediments (Bänziger, 1998), seemed to be synchronised with macro- 
phyte growth. It emerged only once the macrophytes were established, and newly 
hatched larvae seemed to grow fast enabling them to emerge between the end of August 
and September. Thus this species showed a fast summer generation and a slower 
growing winter generation. M. azurea was more abundant on adjacent sediments than 
on macrophytes. It seemed to emerge early and grow slowly, as a limited number of 
larvae from the year emerged until autumn. Fast growth of M. longicornis and slow 
growth of M. azurea were also reported from Petersson (1989) in southern Sweden. 

This study showed that closely related taxa with similar ecological niches may 
coexist in macrophyte beds at some developmental stages. Thus competition seemed 


RUTH BÄNZIGER 


148 


, ‘dds $72220 10 ‘dds sapropisay = Ang usunpos Jusselpe oy) uo pur poq Aydonmeuı AU} JO aus 
ot) UT put 95p9 OU} UT (2292/14/90 ‘O PUL SHISNID] SH2220 ‘SIULOIITUO] JA “Va.inzv Sop1>DISKW) vAAOYOIA, JO SIVISUI FLAIR] JU9I9JJIp Jo Aouanbosy 


+ Old 
pag juejd jo 21)ua9 | ped juejd jo abpa [A lueuipes Juaoelpe El 
saeysul n AI Il Il "Ane A N il Il ane N N HI I ANT: 
7 0) 
4 
4, 
4 
4, os D 
4 o 
7 5 
= 
00, ® 
t6‘ IA SO u 
N N Ml Il ‘Ang A Al Ill Il Ang N N I Il Ang A Al All Il ane À 
F O0 
# 4 5 
dl 7 
4 4 (o) 
2 4 Oy © 
7 4 
g 3 
00L® 
ve HA vO 5 
N N Il I ang N N Il Il ang A N Il [ Ang N N Il I AND m 
d 
7 TE o i 
/ 5 
4, og = 
A 
00! 
9921920 S19930 SIH]SNIE| 5119990 SIU109/BUOJ  S2pl2E]SAW eainze  S9PIE]SAN PEALE 


AQUATIC INSECTS IN MACROPHYTE BEDS 149 


31.V.94 


100 Acentria__ephemerella 


adjacent sediment 
7S edge of plant bed 


F; 
À 50 HB centre of plant bed 7 
È 25 go 
ra 7 
£ 0 | 
i | Il Ill IV V 
5 04.VII.94 
© 100 
= 
© 75 
(cd) 
= 50 
‘© 25 
8 
Ss | Il Ill IV V 
= 05.VIII.94 
A 
© 100 
@ 
> 75 
s 
® 
x 50 


2/8 


| Il Il IV V instars 


Fic. 5 


Frequency of different larval instars of Acentria ephemerella in the edge and in the centre of 
the macrophyte bed and on the adjacent sediment. 


not to be involved in species distribution during most of the invertebrate life cycle. 
Several papers dealing with invertebrate distribution on macrophytes in lakes conclude 
at an absence of competition in this habitat (Magdych, 1979). Hargeby (1990) 
explained this absence of competition by the yearly disturbance undergone by the 
invertebrates through the annual life cycle of the macrophytes. Each year colonization 
has to be resumed and competition has not enough time to take place (Pickett & White, 
1985). 


150 RUTH BÄNZIGER 


Temporal segregation occurred, however, between congeneric species before 
hibernation and during emergence. It may allow better resource partitioning when 
resources are scarce in winter and when maximum energy and food is needed just 
before emergence. Though there was a tendency for some taxa to live on the edge or on 
the adjacent sediment at small size classes and to be more abundant in the centre at the 
last instars or bigger size classes, this (statistically unconfirmed) result may be biased 
by the impact of predation at the edge on large and moving insects. 


ACKNOWLEDGEMENTS 


I am grateful to all friends and colleagues who helped with the field work. In the 
laboratory, I thank B. Oertli for his stimulating discussions and D. Cambin who helped 
with HCW measurements. Special thanks to J.-P. Haenni who provided informations on 
A. ephemerella. | am grateful to B. Lods-Crozet and E. Castella for critical review of 
earlier drafts of this paper. Production of this paper was possible thanks to a grant from 
the Academic Society of Geneva. 


REFERENCES 


BANZIGER, R. 1998. Répartition spatio-temporelle des invertébrés aquatiques en relation avec la 
dynamique des herbiers littoraux (lac Léman). These no 3035, Université de Genève, 
120 pp. + annexes. 

BANZIGER, R., ZANINETTI, L. & LACHAVANNE, J.-B. (submited). Edge effect in aquatic systems: 
macroinvertebrates in lake littoral macrophyte beds. Aquatic Sciences. 

Bass, J. A. B., LADLE, M. & WELTON, J. S. 1982. Larval development and production by the net- 
spinning caddis, Polycentropus flavomaculatus (Pictet) (Trichoptera), in a recirculating 
stream channel. Aquatic Insects 4: 137-151. 

BENGTSSON, B. E. 1981. The growth of some ephemeropteran nymphs during winter in a north 
Swedish river. Aquatic Insects 3: 199-208. 

BENKE, A. C. & JACOBI, D. I. 1986. Growth rates of mayflies in a subtropical river and their 
implications for secondary production. Journal of the North American Benthological 
Society 5: 107-114. 

BERG, K. 1941. Contributions to the biology of the aquatic moth Acentropus niveus (Oliv.). 
Videnskabelige meddelser fra Dansk naturhistorik forening 105: 59-139. 

BRADBEER, P. A. & SAVAGE, A. A. 1980. Some observations on the distribution and life history of 
Caenis robusta Eaton (Ephemeroptera) in Cheshire and North Shropshire, England. 
Hydrobiologia 68: 87-90. 

BRITAIN, J. E. 1982. Biology of Mayflies. Annual Review of Entomology 27: 119-147. 

CLIFFORD, H. F. 1982. Life cycles of mayflies (Ephemeroptera), with special reference to volti- 
nism. Quaestiones Entomologicae 18: 15-90. 

GERKING, S. D. 1957. A method for sampling the littoral macrofauna and its application. Ecology 
38: 219-226. 

HAENNI, J.-P. 1974. Biologie des Pyralidés (Lepidoptera) aquatiques sur la rive sud du lac de 
Neuchatel. Travail de Licence, Institut de Zoologie, Université de Neuchâtel, 108 pp. 

HAENNI, J.-P. 1980. Contribution à la connaissance de la biologie des papillons aquatiques 
(Lepidoptera, Pyraloidea) sur la Rive Sud du Lac de Neuchatel. Bulletin de la Société 
Neuchâteloise de Sciences naturelles 103: 29-43. 

HARGEBY, A. 1990. Macrophyte associated invertebrates and the effect of habitat permanence. 
Oikos 57: 338-346. 


AQUATIC INSECTS IN MACROPHYTE BEDS 151 


HILDREW, A. G. & EDINGTON, J. M. 1979. Factors facilitating the coexistence of hydropsychid 
caddis larvae (Trichoptera) in the same river system. Journal of Animal Ecology 48: 
557-576. 

LANDA, V. 1968. Development cycles of central European Ephemeroptera and their interrelations. 
Acta Entomologica Bohemoslovaca 65: 276-284. 

LEPNEVA, S. G. 1966. Fauna of the USSR. Trichoptera, vol. II, 2. Larvae and pupae of Intergri- 
palpia. ByKHovsKII, B. E. (ed.) Izdatel’stvo “Nauka”, Moskva-Leningrad [Transl. by A. 
Mercado, Prof. O. Theodor for Israel Program for Scientific Translations Ltd.], 700 pp. 

Macan, T. T. 1965. The fauna in the vegetation of a moorland fishpond. Archiv fiir Hydro- 
biologie 61: 18-310. 

Mackay, R. J. & WIGGINs, G. B. 1970. Ecological diversity in Trichoptera. Annual Review of 
Entomology 24: 185-208. 

MAGDYCH, W. P. 1979. The microdistribution of mayflies (Ephemeroptera) in Myriophyllum beds 
in Pennington Creek, Johnston county, Oklahoma. Hydrobiologia 66: 161-175. 

Matas, D. & WALLACE, J. B. 1977. Strategies for coexistence in three species of net-spinning 
caddisflies (Trichoptera) in second-order southern Appalachian streams. Canadian Jour- 
nal of Zoology 55: 1829-1840. 

MCAULIFFE, J. R. 1984. Competition for space, disturbance, and the structure of a benthic stream 

community. Ecology 65: 894-908. 

McGaha, Y. J. 1952. The limnological relations of insects to certain aquatic flowering plants. 

Transactions of the American Miscroscopical Society 355-381. 

MINSHALL, G. W. 1984. Aquatic insect-substratum relationships. Pp. 358-400. In: ROSENBERG, 

V.H. & RESH, D. M. (eds). The ecology of aquatic insects. Praeger Publ., New York. 

MULLER-LIEBENAU, I. 1956. Die Besiedlung der Potamogeton-Zone ostholsteinischer Seen. 

Archiv für Hydrobiologie 52: 470-606. 

MUOTKA, T. 1990. Coexistence in a guild of filter feeding caddis larvae: do different instars act as 
different species? Oecologia 85: 281-292. 

OERTLI, B. 1992. L’influence de trois substrats (Typha, Chara, feuilles mortes) d’un étang 
forestier sur la densité, la biomasse et la production des macroinvertébrés aquatiques. 
These no 2557, Université de Geneve, 283 pp. 

PALMER, C., O’ KEEFE, J., PALMER, A., DUNNE, T. & RADLOFF, S. 1993. Macroinvertebrate func- 
tional feeding groups in the middle and lower reaches of the Buffalo River, Eastern Cape, 
South Africa. I. Dietary variability. Freshwater Biology 29: 441-453. 

PETERSSON, E. 1989. Swarming activity patterns and seasonal decline in adult size in some caddis 
flies (Trichoptera: Leptoceridae). Aquatic Insects 11: 17-37. 

PICKETT, S. T. A. & WHITE, P. S. 1985. The ecology of natural disturbances and patch dynamics. 
Academic press Inc. Ltd., London, 472 pp. 

RESH, V. H., 1979. Sampling variability and life history features: basic considerations in the 
design of aquatic insect studies. Journal of the Fisheries Research Board of Canada 36: 
290-311. 

Rooke, J. B. 1984. The invertebrate fauna of four macrophytes in a lotic system. Freshwater 
Biology 14: 507-513. 

Smock, L. A. 1980. Relationships between body size and biomass of aquatic insects. Freshwater 
Biology 10: 375-383. 

SWEENEY, B. W. & VANNOTE, R. L. 1981. Ephemerella mayflies of White Clay Creek: bioener- 
getics and ecological relationships among six coexisting species. Ecology 62: 1353-1369. 

TUDORANCEA, C. & GREEN, R. H. 1975. Distribution and seasonal variation of benthic fauna in 
lake Manitoba. Verhandlungen der Internationalen Vereinigung fiir Limnologie 19: 
616-623. 

WINTERBOURN, M. J. 1971. The life-history and trophic relationships of the Trichoptera of Marion 
Lake, British Columbia. Canadian Journal of Zoology 49: 623-635. 


N 


SEEN, 


ah rs if 
| LUE Pc Fe 
i nds: L' 


ve, La ee 
et M 


MT: 


FR = sea 
è ea Ld Me 


REVUE SUISSE DE ZOOLOGIE 107 (1): 153-163; mars 2000 


Les Nanophthalmus Motschulsky d’Europe 
(Coleoptera, Scydmaenidae) 


Claude BESUCHET! & Stanislav VIT? 
| Muséum d’histoire naturelle, case postale 6434, CH-1211 Genève 6. 
2 Rue de la Poterie 26, CH-1202 Genève. 


European species of the genus Nanopthalmus Motschulsky (Coleoptera, 
Scydmaenidae). - Two new species of the genus Nanophthalmus Motschul- 
sky, 1851, N. nonveilleri sp. n. et N. serbicus sp. n. are described from Yugo- 
slavia (Serbia), N. ditomus Saulcy, treated previously as synonym of N. 
megaloderoides Motschulsky is revalidated and the lectotype of the species 
designated. Additional data related to N. megaloderoides Motschulsky, 
N. turcicus Reitter and N. beszedesi Reitter are provided together with a key 
to all treated species. 


Key words: Coleoptera - Scydmaenidae - Cephenniini - Nanophthalmus - 
taxonomy - Europe. 


INTRODUCTION 


Seulement sept especes ont été décrites dans le genre Nanophthalmus Mot- 
schulsky, 1851: megaloderoides Motschulsky, 1851 de Crimée (espece-type); ditomus 
(Saulcy, 1878) du Caucase; rotundicollis (Reitter, 1881) de Talysch; armeniacus 
Reitter, 1884 du Caucase: turcicus Reitter, 1894 de Turquie d'Europe; robustus Roubal, 
1913 du Caucase et beszedesi Reitter, 1913 de l’Istrie. Mais plusieurs espèces nouvelles 
ont été découvertes au cours des vingt dernières années, par le Professeur Guido 
Nonveiller en Serbie, par M. Rudolf Rous dans le Caucase, par M. Antoine Senglet 
dans le nord de l'Iran, et enfin par nous en Turquie d’Asie. Des lacunes importantes 
sont ainsi comblées dans l’aire de répartition du genre Nanophthalmus. 

Nous avons commencé la révision de ce genre, devenue indispensable. Mais 
l’échéance du “Catalogue of the Palearctic Coleoptera”, nous contraint à une première 
mise au point, à savoir la redescription des Nanophthalmus megaloderoides Mot- 
schulsky, turcicus Reitter et beszedesi Reitter, la revalidation de N. ditomus Saulcy et la 
description de deux espèces nouvelles de Serbie. Ces six Nanophthalmus appartiennent 
au groupe d’espèces de petite taille, ne dépassant en general pas 1 mm. 


Manuscrit accepté le 09.02.1999 


154 CLAUDE BESUCHET & STANISLAV VIT 


MATERIEL ET METHODES 


Abréviations utilisées: 


HNHM Hungarian Natural History Museum, Budapest, Hongrie 

IPVB Institut pour la Protection des Végétaux, Belgrade, Yougoslavie 

MHNG Museum d’histoire naturelle, Geneve, Suisse 

MNHN Museum National d’Histoire naturelle, Paris, France 

MCSN Museo Civico di Storia Naturale “Giacomo Doria”, Gênes, Italie 

MSNT Museo Civico di Storia Naturale, Trieste, Italie 

NHMW Naturhistorisches Museum, Wien, Autriche 

SMNS Staatliches Museum für Naturkunde, Stuttgart, Allemagne 

SMFD Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt/Main, 
Allemagne 

ZMHB Museum fiir Naturkunde der Humboldt Universitat, Berlin, Allemagne 


Autres abréviations: A. - antennes; E. - élytres; Lo. - longueur; La. - largeur: P. - 
pronotum; les étiquettes des matériaux étudiés sont citées ici de maniere strictement 
conforme a l’original:/.../.- étiquette citée: /”....”/ - passages manuscrits de I’ étiquette. 


DESCRIPTIONS 
Genre Nanophthalmus Motschulsky, 1851 Biesse? 


Nanophthalmus Motschulsky, 1851: 506; espèce-type Nanophthalmus megaloderoides Mot- 
schulsky, 1851, par monotypie. 

Cephennium (Nanophthalmus), Reitter, 1881: 554 

Nanophthalmus, Cziki, 1919:17 

Nanophthalmus, Lazorko, 1962: 277 

Nanophthalmus, Newton & Franz, 1998: 143 


DIAGNOSE. Scydmaenidae Cephenniini anophtalmes et aptères. Téte petite, les tempes 
saillantes. Antennes (Fig. I) de 11 articles, la massue bien développée, formée de deux 
articles seulement. Pronotum nettement convexe, un peu moins large que les élytres; 
côtés arrondis, finement rebordés; aucune ornementation particulière sur le pronotum; 
pas d’apophyse prosternale. Elytres convexes, sans carène laterale, ornés chacun d’une 
petite fossette basale profonde, tomenteuse, accompagnée du còté externe d’une caré- 
nule très courte; pas de carène humérale à proprement parler mais un pli normalement 
caché sous la base du pronotum; apex des élytres souvent rebordé. Edéage formé d’une 
grande capsule basale plus ou moins nettement prolongée par une sorte de manchon 
dans lequel se trouve l’armure copulatrice; paramères gréles, atteignant l’apex de 
l’édéage, terminés par une soie. 


Nanophthalmus megaloderoides Motschulsky, 1851 Fe 


Nanophthalmus megaloderoides Motschulsky, 1851: 506 
Nanophthalmus megaloderoides; Motschulsky, 1869: 271; 
Cephennium (Nanophthalmus) megaloderoides; Reitter, 1881: 555; 
Cephennium (Nanophthalmus) megaloderoides; Reitter, 1884:84; 
Nanophthalmus megaloderoides: Lazorko, 1962: 314-316, fig. 15; 


LES NANOPHTHALMUS MOTSCHULSKY D'EUROPE 155 


MATERIAUX EXAMINES: Type perdu. (Loc.typ. Ukraine, Aloupka). 31 ex. /laila Gebirge, Krim, 
Winkler/ (HNHM, MCSN, MHNG, MSNT, NHMW, SMFD, ZMHB). 

La collection Victor de Motschulsky du Muséum de Moscou ne renferme aucun 
exemplaire du genre Nanophthalmus et l’exemplaire ayant servi à la description origi- 
nale semble perdu. L’espece a été décrite de la Crimée (Tauride, Aloupka). Reitter 
(1881) n’a pas dü voir des exemplaires de Motschulsky comme le montre sa description 
qui n° est en fait qu'une mise en commun des données publiées par Motschulsky et de 
Saulcy (ditomus). La diagnose ultérieure et l’édéage publiés pour l’espece megalo- 
deroides Motschulsky par Lazorko (1962) s’appuient sur des exemplaires provenant des 
Monts Aj Petri, pris par Moczarski et Winkler en mai 1911, qui figurent dans diverses 
collections, étiquetés collectivement laila-Gebirge Krim, Winkler (ou) Moczarski. Ces 
exemplaires concordent avec la description originale et appartiennent a une seule 
espece, bien caractérisée par son édéage, la seule du genre Nanopththalmus connue jus- 
qu'à present de la Crimée. C’est l’espèce la plus occidentale des territoires septen- 
trionaux de la Mer Noire. 


DIAGNOSE. Longueur (tête comprise) 0,87-0,97 mm. Elytres moins allongés que chez 
les espèces balcaniques, ratio E.Lo./E.La. = 1,34-1,41; extrémités apicales des élytres 
faiblement modifiées, bords épaissis sans être nettement relevés. Pronotum sur la ligne 
médiane à peu près aussi long que large à la base; côtés du pronotum arqués, 
convergeant en arrière vers les angles postérieurs; base droite, ratio P.Lo./P.La. = 0,78- 
0,88; largeur du pronotum subégale ou inférieure à la longueur des antennes; dépres- 
sions basales des élytres effacées: angles huméraux avec une fine carène prolongée en 
arrière. 

Antennes: ratio A.Lo./ P.base = 1,14-1,34; articles 3-8 courts, suboblongs ou 
sphériques, article 9 petit et subsphérique chez les males, plus grand, pyriforme et plus 
transverse chez les femelles; article 11 allonge. 

Face ventrale: carénules du métasternum courtes et écartées. 

Caracteres sexuels des males. Protibias recourbés et plus brusquement renflés 
dans la moitié apicale; face ventrale de la portion basale du profémur pourvue d’une 
fine carene terminée distalement par une minuscule saillie obtuse; métasternum ample- 
ment déprimé (aplati chez la femelle). 

Edéage (Fig. 3). 


DISTRIBUTION. Ukraine, Crimée méridionale: Iaila Gebirge = Monts Aj Petri (nom 
actuel). Aloupka = Alupka, la localité typique, se trouve sur le versant sud de cette 
chaine monagneuse. 


Nanophthalmus ditomus (Saulcy, 1878) Fig. 4 


Cephennium ditomum Saulcy, 1878: 139-140, pl. II, fig. 21 

Cephennium (Nanophthalmus) megaloderoides Motschulsky = ditomum Saulcy; Reitter, 1881: 
555 

Cephennium ditomum Saulcy = ? Cephennium (Nanophthalmus) megaloderoides Motschulsky; 
Lazorko, 1962: 316 

MATERIEL ETUDIE. Lectotype (par présente désignation) d : (sans provenance), /disque or/, /”dito- 

mum n. sp. Saulcy, Typ.’/,/" 14” (étiquette bleue et noire)/, /Typus (étiquette rouge)/, MUSEUM 


156 CLAUDE BESUCHET & STANISLAV VIT 


DE PARIS 1900 coll. J. CROISSANDEAU/. (MHNP); 14, 12, /disque or/, /disque bleu/ 
” Caucase” (étiquette rose)/, /"Cephenn. ditomum Slcy - Cauc (Rtt)”/, /”14” (étiquette bleue et 
noire)/, /(étiquette rouge vide)/, MUSEUM DE PARIS 1900 coll. J. CROISSANDEAU/; 16, 
/disque or/, /Caucasus, Meskisches Geb. Leder, (Reitter)/, /’Cephenn. ditomum Saulcy, 
Caucas”/,/"29” (étiquette bleue et noire)/, /(étiquette bleue)/, MUSEUM DE PARIS 1900 coll. J. 
CROISSANDEAU/; 1d, /disque or/, /Caucasus, Meskisches Geb. Leder, (Reitter)/, /°14” 
(étiquette bleue et noire)/, MUSEUM DE PARIS 1900 coll. J. CROISSANDEAU/ (tous les 
exemplaires MNHN). Autres:8¢d, 1222, /Caucasus, Meskisches Geb. Leder (Reitter)/, 
(MNHN, MHNG, NHMW, HNHM, MCSN). 

Nanophthalmus ditomus Saulcy est une bonne espece. Nous avons pu examiner 
des matériaux de l’époque, retrouvés dans la collection Croissandeau, et constater que 
la mise en synonymie de cette espece par Reitter (1881:555) n’était pas fondée. Faute 
de matériel disponible de Suram, localité-type de ditomus, Lazorko (1962:316) main- 
tient formellement la synonymie de Reitter tout en soulignant son caractere douteux. 
Nous avons choisi pour lectotype un male de la collection Croissandeau du Muséum de 
Paris, portant l’étiquette manuscrite de Saulcy (voir ci-dessous). 


DIAGNOSE. Longueur (téte comprise) 0,92-0,99 mm (Saulcy 0.75 mm!). Elytres moins 
allongés, ratio E.Lo./E.La. = 1,35-1,45; extremites apicales des élytres modifiées, bords 
faiblement rebordés et relevés. Pronotum sur la ligne médiane plus court que large a la 
base; côtés du pronotum arqués, convergeant en arrière vers les angles postérieurs; base 
droite, ratio P.Lo./P.La. = 0,81-0,86; largeur du pronotum généralement légèrement 
supérieure ou tout au plus égale a la longueur des antennes; dépressions basales des 
élytres pratiquement indistinctes; angles huméraux tout au plus avec une tres courte 
carene. 

Antennes distinctement plus longues que le pronotum sur la ligne médiane ou sa 
largeur basale, ratio A.Lo./P.base = 1,05-1,19; articles 3,4 et 5 suboblongs, 6,7 et 8 
courts, subsphériques; article 9 généralement assez gros, subpyriforme et transverse 
chez le deux sexes; article 11 un peu plus long que large. 

Face ventrale: carénules du métasternum courtes et écartées. 

Caracteres sexuels des males. Protibias faiblement recourbés, apex avec une 
protubérance dentiforme; face ventrale de la portion basale du profémur avec une fine 
carene s’estompant distalement; métasternum amplement concave (faiblement convexe 
chez la femelle). 

Edéage (Fig. 4). 
DISTRIBUTION. Géorgie, Suram (localité-type), situé à l'extrême nord-est de Meshedskiy 
Khrebet (= Meskisches Gebiet des auteurs allemands). 


Nanophthalmus turcicus Reitter, 1894 His 


Nanophthalmus turcicus Reitter, 1894: 114. 


MATERIEL ETUDIE. Holotype d étiquetté:/’Nanophthal. turcicus m. 1894”/, /’Eur. Turkei 
Merkl.”/, /disque or/, MUSEUM DE PARIS 1900 coll. J. CROISSANDEAU/. (MNHN) 
/Typus/”tureicus Rtt.” Cl. Besuchet det. XII. 1957/. (MHNP); 26 8, 12 / “Belgrader Wald près 
Istanbul, 8.V.58, H. Schweiger”/. (MHNG). 


DIAGNOSE. Longueur (tete comprise) 0,94-1,05 mm (Reitter 0,8 mm!). Elytres moins 
allongés, ratio E.Lo/E.La. = 1,28-1,38; extrémités apicales des élytres a peine 


LES NANOPHTHALMUS MOTSCHULSKY D’EUROPE 157 


Fics 1-4 


l. Nanophthalmus sp., antenne; 2. Nanophthalmus sp., apex du protibia; 3. Nanophthalmus 
megaloderoides Motschulsky, édéage face ventrale; 4. Nanophthaimus ditomus (Saulcy), édéage 
face ventrale. (Echelle en mm). 


158 CLAUDE BESUCHET & STANISLAV VIT 


modifiées, simplement renflées. Pronotum sur la ligne médiane aussi long que large à la 
base; cotés du pronotum arqués, convergeant en arriere vers les angles postérieurs; base 
droite, ratio P.Lo./P.La. = 07,8-0,82; largeur du pronotum légèrement inférieure à la 
longueur des antennes; dépressions basales des élytres pratiquement indistinctes; angles 
huméraux tout au plus avec une très courte carène. 

Antennes très nettement plus longues que le pronotum sur la ligne médiane , 
ratio A.Lo./P. base = 1,13-1,28; articles 3 et 4 subégaux, oblongs, 7 et 8 courts, 
subsphériques; article 9 plus gros, transverse chez le deux sexes; article 11 moins d’une 
fois et demie plus long que large. 

Face ventrale: carénules du métasternum courtes mais peu écartées. 

Caracteres sexuels des males: protibias faiblement recourbés et renflés dans leur 
moitié apicale; crochet apico-interne petit, peu distinct; face ventrale de la portion 
basale du profémur pourvue d’une fine carene terminée distalement par une minuscule 
saillie obtuse; métasternum amplement concave chez le male, simplement applati chez 
la femelle. 

Edéage (Fig. 5). 


DISTRIBUTION. Turquie, Thrace, Forét de Belgrade pres d’Istanbul. 


Nanophthalmus beszedesi Reitter, 1913 Fig. 6 
Nanophthalmus beszedesi Reitter, 1913: 140-141. 

MATERIAUX EXAMINES. Lectotype (par présente désignation) 9: étiqueté, /Dr. v. Beszédes, M. 
Maggiore, Istrien, “15/8°/ Mus. HMHN det. Reitter Nanophthalmus beszedesi. Paralectotype (par 
présente désignation) 1 ©, /idem/, (les deux HNHM). Il s’agit bien des matériaux vus par Reitter 
qui cite “2 ex. am 15 August v. J.”. Autres: 1¢, /Dr. Beszedes, M. Maggiore, Istria “22.V. 
1913”/, /’Nanophthalmus Beszedesi Reitter”/. (MHNG); 14, /idem/. (MCSN.- coll Dodero); 14, 
/Mte Maggiore, Istr. Winkler. (ZMHB); 3 ex. /Mte Maggiore, Winkler/. (SMFD). 

DIAGNOSE. Longueur (tête comprise) 0,87-0,92 mm. Elytres très allongés, ratio E.Lo./ 
E.La. = 1,48-1,53; extrémités apicales des élytres tres nettement modifiees, présentant 
un fort bourrelet retroussé vers le haut, échancré a la suture. Pronotum sur la ligne 
médiane aussi long que large a la base: côtés du pronotum fortement arqués au tiers 
antérieur, subparallele en arriere avant les angles postérieurs; ratio P.Lo./P.La. = 0,87- 
0,88; largeur du pronotum égale ou inférieure à la longueur des antennes; callosités 
humérales distinctes; dépressions basales indistinctes. 

Antennes nettement plus longues que le pronotum sur la ligne médiane ou 
comme sa largeur à la base, ratio A.L/ P.base = 1,14-1,2 ; articles 3, 4 et 5 à peine 
oblongs, 6, 7 et 8 subsphériques, article 9 petit, subsphérique chez le mâle, plus grand et 
pyriforme chez la femelle: article 11 court, tout au plus d’un quart plus long que large. 
Face ventrale: carénules mésosternales prolongeant la lame sternale à peine déve- 
loppées. 

Caractères sexuels des mâles. Protibias recourbes, brusquement renflés dès le 
milieu, apex avec une protubérance dentiforme; face ventrale de la portion basale du 
profémur sans caractères particuliers; métasternum amplement déprimé (aplati chez la 
femelle). 

Edéage (Fig. 6). 
DISTRIBUTION. Croatie, Istrie, Monte UËka, (= Monte Maggiore). 


LES NANOPHTHALMUS MOTSCHULSKY D'EUROPE 159 


Nanophthalmus nonveilleri sp. n. Figs 78, 11 


MATERIAUX EXAMINES. Holotype. d, étiqueté: /Serb. Tara pl., 1000 m, “11.06. 84”, Nonvll./ 
(IPVB); Paratypes. 14, 29 2, / idem “11.06.84”/. (MHNG); 28 d, 29 2,./ idem, “23.05. 84”/ 
(IPVB, MHNG); 16, / idem, “6.10. 78”/ (IPVB); 1d, /Serb. Tara pl., 900 m, “6.10. 78/ (IPVB). 
16 /Serb. Tara pl., 900 m, “6.10.78/ (IPVB). 14, 29 2, /Yougoslavie, Serbie: Tara Planina, 
Mitrovac (route Mitrovac-lac d’accumulation), 900 m, 23.V.1984; tamisage de feuilles mortes 
dans une forêt de hétres, érables et épicéas (Besuchet)/. (MHNG). 

DIAGNOSE. Longueur (tête comprise) 0,89-1,02 mm. Elytres très allongés, ratio Long./ 
Larg. = 1,43-1,55; extrémités apicales des élytres non modifiées, simples. Pronotum sur 
la ligne médiane plus long que large à la base; côtés du pronotum fortement arqués au 
milieu, convergeant en arrière nettement vers les angles postérieurs; base droite; ratio 
P.Long./P.Larg. = 0,86-0,91; largeur du pronotum supérieure a la longueur des an- 
tennes; base des élytres sans trace de dépression basale ou de callosité humérale. 

Antennes un peu ou nettement plus longues que le pronotum sur la ligne 
médiane et tres nettement plus longues que la largeur basale de celui-ci, ratio A.L/ 
P.base = 1,09-1,34; articles 3, 4 et 5 oblongs, 6, 7 et 8 subsphériques, article 9 trans- 
verse, pyriforme, plus nettement transverse semble-t-il chez les males et plus allonge 
chez les femelles: article 11 court, un peu plus long que large. 

Face ventrale: carénules du métasternum faiblement divergentes. 

Caracteres sexuels des males. Protibias recourbés, nettement renflés dans la 
moitié distale, pourvus de crochet apical; face ventrale de la portion basale du profemur 
avec une fine carene s’estompant distalement; metasternum peu differencie, a peine 
aplani chez le male (faiblement convexe chez la femelle). 

Edéage. (Figs. 7, 8). 


DISTRIBUTION (Fig. 11). Serbie occidentale:Tara planina ( (8) localité-type). Selon Dr. 
G. Nonveiller (communication personnelle) cette espece se rencontre aussi a Goc 
planina, Beli izvori et Dobre vode (9); Ivanjıca, Mucina voda (10); Ovcar Banya, 
Ovcarsko-Kablarska klisura, Debela Gora (11). Elle va de 600 a 1000 m d’altitude. 
(Ces exemplaires ont été capturés et determinés par MM. Nonveiller et Pavicevic et 
sont déposés a IPVB). 


Nanophthalmus serbicus sp. n. Bios) 910-3 


MATERIAUX EXAMINES. Holotype: d, étiqueté /Serb. Stol pl., 800 m, Nonvll., “26.05. 198477. 
(IPVB). Paratypes: 16, /idem “26.05.1984”/. (MHNG); 2 ex. /idem ‘ ‘10.04.1987 1 (IPVB, 
MNHG); 38 d, 22 9, /idem “15.5.1982” (IPVB, MHNG); 14 /Serb. Rtanj pl. “12.III. 1977.”/ 
(MHNG): 96 d,79 2, / Yougoslavie, Serbie: Stol Planina (30 km au nord de Bor), 900 m, 26.V. 
1984; tamisage de feuilles mortes au pied de rochers, dans une forét de hétres, (Besuchet). 
(MHNG); Tel 39 2,/ même provenance, 27.V.1984; lavage de terre au pied de vieilles souches 
de hétres (Besuchet). (MHNG); 26 8, 12, /Yougoslavie, Serbie: Rtanj Planina, 600 m, 27.V. 
1984: tamisage de feuilles mortes au fond d’un ravin, dans la forét de hétres (Besuchet). 
(MHNG). 


DIAGNOSE. Longueur (téte comprise) 0, 92-0,99 mm. Eytres nettement allongés, ratio 
Long./Larg. = 1,39-1,5; extrémités apicales des élytres a peine modifiées, bords épaissis 
sans €tre nettement relevés. Pronotum sur la ligne médiane plus court ou tout au plus 
aussi long que large à la base; côtés du pronotum fortement arqués dans le tiers anté- 


160 CLAUDE BESUCHET & STANISLAV VIT 


Figs 5-10. Nanophthalmus, édéage face ventrale - 5. Nanophthalmus turcicus Reitter, (Holo- 
type); 6. Nanophthalmus beszedesi Reitter; 7. Nanophthalmus nonveilleri sp.n., (Paratype), 8. 
variabilité de l’apex de l’édéage (Paratype); 9. Nanophthalmus serbicus sp.n. (Paratype, Rtanj 
planina), 10. variabilité de l’apex de l’édéage (Paratype, Stol planina). (Echelle en mm). 


LES NANOPHTHALMUS MOTSCHULSKY D’ EUROPE 161 


HONGRIE 


e 
Subotica 


ROUMANIE 
Novi Sad 
© 
Beograd 
® 
NEE Waliewo SERBIE I Ai 
® 
1 
LA 
Uzice Kragujevac Zajecar 
8 e 
x 11 2 
9 3 BULGARIE 
10 @ Krusevac 
6 
e 
Nis 
e 
CROATIE Pristina 
Podgorica 
Jy ° 
ni 
e Skopje 
ALBANIE 
MACEDOINE 
\ 
Fic. 11 


Carte de distribution - espèces de la Serbie: Nanophthalmus serbicus sp. n. (No. 1 - 7); Nano- 
phthalmus nonveilleri sp. n. (No. 8 - 11). 


162 CLAUDE BESUCHET & STANISLAV VIT 


rieur, presque subparalleles avant les angles apicaux, base droite, ratio P.Long./ P.Larg. 
= 0.81-0,86; largeur du pronotum subégale ou légèrement inférieure a la longueur des 
antennes; angles huméraux oblitérés, nuls; dépressions basales des élytres faibles. 

Antennes tres nettement plus longues que le pronotum sur la ligne médiane et 
nettement plus longues que sa largeur basale, ratio A.L/ P.base = 1,08-1,33; articles 3, 4 
et 5 oblongs, 6, 7 et 8 subsphériques, article 9 petit, subsphérique à faiblement trans- 
verse chez les males, plus grand, pyriforme et plus nettement transverse chez les 
femelles, article 11 court, pratiquement aussi large que long. 

Face ventrale: carénules du métasternum fortement divergentes. 

Caracteres sexuels des males. Protibias plus nettement recourbés dans leur 
portion apicale seulement, nettement renflés des le milieu, dépourvus d’un crochet 
apical; face ventrale de la portion basale du profémur avec une fine carene s’estompant 
distalement; métasternum fortement déprimé, concave (applati chez la femelle,). 
Edéage (Figs. 9, 10). 


DISTRIBUTION (Fig 11). Serbie orientale:(1)Stol planina, (localité-type) et (2)Rtanj 
planina;. Selon Dr. G. Nonveiller (communication personnelle) cette espece se ren- 
contre aussi à Zajecar- Lenovac (3); Homolje - Ceremosnja (4); Mts. Homolje - 
Mejdan Kucana (5); Jastrebac planina - Ravniste (6) et Kragujevac - Zezelj (7). Elle va 
de 450 à 1200 m d altitude. Ces exemplaires ont été capturés et determines par MM. 
Nonveiller et Pavicevic (déposés a IPVB). 


TABLEAU DES ESPECES CITEES 


Ce tableau n’a qu'une valeur indicative; les identifications doivent être contrôlées par 
l’édéage. 


| Apex des élytres orné d’un bourrelet fortement développé, relevé vers le 
HAMSTER ER eae Vis wed os os E beszedesi Reitter 
Apex des élytres non relevé, tout au plus avec un bourrelet léger........... 2 
2 Elytres plus larges (0,46-0,49 mm), tres convexes; Thrace..... turcicus Reitter 
: Elytres moins larges (0,40-0,43 mm), moins convexes .................... 3 
3 Pubescence du pronotum et des élytres formée de soies couchées de 
longueur moyenne; élytres plus courts (indice élytral 1,35-1,41)............ - 
- Pubescence du pronotum et des élytres formée de soies semi-dressées 
assez longues: élytres plus allongés (indice élytral 1,42-1,55).............. 5 
4 Corps moins déprimé (haut. - 0,35 mm): en vue latérale la convexité du 
pronotum est nettement séparée de celle des élytres; Géorgie . ditomus (Saulcy) 
- Corps relativement déprimé (haut. ~ 0,30 mm): en vue latérale la 


convexité du pronotum se prolonge sur celle des élytres; Crimée 

A Rael Nope Varn SARE URI megaloderoides Motschulsky 
5) Antennes légèrement plus longues, les articles 6 et 7 légèrement plus 

longs que larges; métasternum du mâle avec une dépression arrondie 

profonde3Serbie tr. 1 LIETA Oe ee ee ee serbicus sp. n. 
- Antennes légèrement plus courtes, les articles 6 et 7 à peine ou lége- 

rement plus larges que longs; métasternum du male avec une 

dépression triangulaire très superficielle; Serbie............. nonveilleri Sp.n. 


LES NANOPHTHALMUS MOTSCHULSKY D’ EUROPE 163 


REMERCIEMENTS 


Nous tenons à remercier ici toutes les institutions citées, pour nous avoir facilité 
l'étude des matériaux des collections historiques, traités dans cette note, puis plus 
particulièrement le Professeur Guido Nonveiller de Zemun et l’Institut pour la 
Protection des Végétaux de Belgrade, Yougoslavie, pour nous avoir confié l’étude des 
matériaux provenant de leurs recherches. 


BIBLIOGRAPHIE 


CSIKI, E. 1919. Scydmaenidae. /n:Schenkling, S. (ed.). Coleopterorum Catalogus, Pars 70. Berlin, 
W. Junk, 106 pp. 

LAZORKO, W. 1962. Zwei neue Cephennium-Arten (Col. Scydmaenidae) mit einer Übersicht der 
ukrainischen Arten der Tribus Cephenniini. Entomologische Arbeiten aus dem Museum 
G. Frey, Tutzing 13(2): 273-320. 

MOTSCHULSKY, V. DE. 1851. Enumération des nouvelles espèces de Coléoptères II. Bulletin de la 
Societe Imperiale des Naturalistes de Moscou 24(2): 479-511. 

MOTSCHULSKY, V. DE. 1869. Enumération des nouvelles espèces de Coléoptères rapportés de ses 
voyages (7e), Scydmaenides. Bulletin de la Société Impériale des Naturalistes de Moscou 
42(1): 252-272. 

NEWTON, A. F. & FRANZ, H. 1998. World catalogue of the genera of Scydmaenidae (Coleoptera). 
Koleopterologische Rundschau 68: 137-165. 


REITTER, E. 1881. Bestimmungs-Tabellen der europäischen Coleopteren. V. Paussidae, Clavi- 
geridae, Pselaphidae und Scydmaenidae. Verhandlungen der Kaiserlich-Königlichen 
Zoologisch-Botanischen Gesellschaft in Wien 31: 443-593, pl. 6, 7. 

REITTER, E. 1884. Bestimmungs-Tabellen der europäische Coleopteren. X. Nachtrag zu den V. 
Theile, enthaltend:Clavigeridae, Pselaphidae und Scydmaenidae. Verhandlungen der 
Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 34: 59-94. 

REITTER, E. 1894. Neue Pselaphiden und Scydmaeniden aus der europäischen Türkei. Wiener 
Entomologische Zeitung 13: 113-115. 

REITTER, E. 1913. Eine Serie neuer Scydmaeniden aus der europäischen Fauna. Entomologische 
Blätter 9: 139-143. 

ROUBAL J. 1913. Zwei neue paläarktische Coleopteren. Entomologische Mitteilungen, Berlin- 
Dahlem 11(1): 21-22. 

SAULCY, F. DE. 1878. (Pselaphidae, Scydmaenidae). /n: Schneider, O. & Leder, H. Beiträge zur 
Kenntniss der kaukasischen Käferfauna. Verhandlungen des Naturforschenden Vereines 
in Brünn (1877) 16: 3-258, pls. 1-4. 


li Va j 
et 


| MGR Yes 


Tate 


REVUE SUISSE DE ZOOLOGIE 107 (1): 165-211; mars 2000 


Über die Lauterzeugung der Welse 
(Siluroidei, Ostariophysi, Teleostei) und ihren Zusammenhang 
mit der Phylogenie und der Schreckreaktion * 


Andreas HEYD & Wolfgang PFEIFFER 
Zoologisches Institut, Universität Tübingen, 
Auf der Morgenstelle 28, 

D-72076 Tübingen, Germany. 


Sound production in catfish (Siluroidei, Ostariophysi, Teleostei) and its 
relationship to phylogeny and fright reaction. - Sound production by 
pectoral stridulation, and sound production by swim-bladder mechanisms 
were studied in 19 species of catfish from 8 families. Stridulation sounds 
were recorded in air by means of a microphone and a tape recorder while 
holding one of the pectoral spines; swim-bladder sounds were recorded 
under water. All sounds were investigated by sonographic analysis of the 
records. In addition, three species (Arius seemani, Pimelodella gracilis, 
Pimelodus pictus) were studied anatomically in order to determine whether 


or not they have a swim-bladder mechanism. 


Sound production by stridulation was observed when the enlarged pectoral 
fin spines were moved in both directions (spread and attracted) in Mystus 
gulio, M. vittatus (Bagridae), Synodontis ocellifer, S. schoutedeni (Mo- 
chokidae), Agamyxis flavopictus (Doradidae), and Dysichthys coracoideus 
(Aspredinidae). In contrast, the following species produced sounds only 
when they spread their pectorals: Arius seemani (Ariidae), Pimelodella (= 
Brachyrhamdia) meesi (Pimelodidae), Dianema urostriata (Callichthy- 
idae), Peckoltia pulcher, and Glyptoperichthys gibbiceps (Loricariidae). In 


the other Loricariidae studied sound production was absent. 


In addition to the pectoral stridulation mechanism, Arius seemani possesses 
a modified swim-bladder mechanism, called the "Springfederapparat“ 
(Müller, 1842) or „elastic spring apparatus“. Thin elastic bones, derived 
from the transverse processes of the first few vertebrae, function in sound 
production. Specialized sonic muscles on the upper surface of this „elastic 
spring“ cause the vibration of the swim-bladder. The elastic bones function 
as antagonists of the muscular contraction. They return the fibres of the 
sonic muscles after each contraction. The sonic activity of A. seemani with 
its swim-bladder mechanism showed a daily rhythm. The choruses were 
most vigorous in the early morning. The sounds emitted consisted of short 


* In memoriam Prof. Dr. Karl von Frisch 
Manuskript angenommen am 9.11.1999 


166 


ANDREAS HEYD & WOLFGANG PFEIFFER 


pulses at irregular intervals with a fundamental frequency near 170 + 30 
Hz. This frequency was equivalent to the vibration frequency of the sonic 
muscles. 

In Pimelodus pictus (Pimelodidae) the protractor muscle (functioning as a 
vibration generator) is directly connected to the swim-bladder (serving as a 
resonance body). This system generates drumming sounds. In Pimelodella 
gracilis a swim-bladder mechanism could not be found. 

We detected a new method of sound production in Hemibagrus nemurus 
(Bagridae). It is able to squeak or to scream by pressing out air from its gill 
slits. 

The frequent occurence of a stridulatory mechanism by means of the 
pectoral spines in at least 13 out of 33 families of catfish indicates that 
those species which lack a stridulatory mechanism may have subsequently 
lost it during evolution. In contrast, swim-bladder mechanisms are 
supposed to have developed independently several times, since they are 
differently structured. The existence of two very different sonic mecha- 
nisms, a stridulatory as well as a swim-bladder mechanism, in the same 
species (for example Arius seemani and Pimelodus pictus) indicates that 
the sounds may have a different biological significance. The frequencies of 
the nonharmonic sounds by stridulation (described as chirps, clicks, croaks, 
grunts, knocks, scrapes, scratches, squeaks, rasps, thumps and so on) range 
from 100 - 8000 Hz, mostly 1000 - 4000 Hz. Their main acoustic energy is 
found at frequencies above the hearing range of most fish, except the 
Ostariophysi. In addition, all predaceous fishing mammals and birds are 
able to hear these frequencies. Thus, these sounds may serve as an impor- 
tant means of warning and defence, especially since they are mostly 
produced when the catfish is captured and pulled out of the water. The 
noise may function in the same way as that produced by threatened rattle 
snakes. In a similar manner to these poisonous snakes, the stridulating 
catfish are defensive. The pectoral spines and the dorsal spine are erected 
simultaneously in order to protect the fish. Since pectoral and dorsal spines 
have locking mechanisms, predators can swallow these catfish only with 
great difficulty. Whereas stridulatory mechanisms exist in many solitary 
species, swim-bladder mechanisms are numerous in social catfish and are 
supposed to serve intraspecific communication. 

In all species of the mostly solitary and night active catfish that have been 
studied with respect to their sound production, the fright reaction elicited 
by the alarm substance from the epidermis of conspecifics is either little 
developed (Mochokidae, Pimelodidae) or absent (Aspredinidae, Lori- 
cariidae). On the other hand, no stridulatory mechanism has been detected 
in some pelagic, schooling species of catfish possessing a distinct, well- 
developed fright reaction and being active in daylight (Eutropiellus van- 
deweyeri - Schilbeidae, Kryptopterus bicirrhis - Siluridae). However, since 
only a few species of catfish have been studied, conclusions concerning a 
connection between the existence of the chemically elicited fright reaction 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 167 


on the one hand and sound production by sonic mechanisms on the other 
should only be drawn with caution. 


Key-words: fish - catfish - sound production - stridulation - swim-bladder 


mechanisms (Trommelmechanismen) - „elastic spring apparatus“ 
(Springfederapparat) - fright reaction (Schreckreaktion) - alarm substance 
(Schreckstoff). 

EINLEITUNG 


Die Welse (Siluroidei) umfassen mit etwa 2500 rezenten Species in 33 Fami- 
lien (Teugels, 1996) mehr als ein Drittel aller Ostariophysen (Sagemehl, 1885) und 
ungefähr 10% der Teleostei (Eigenmann & Eigenmann, 1890; Regan, 1911; Berg, 
1958; Greenwood er al., 1966; Roberts, 1973; Gosline, 1975; Novacek & Marshall, 
1976; Briggs, 1979; Fink & Fink, 1981, 1996; Lauder & Liem, 1983; Lundberg & 
McDade, 1986: Nelson, 1994). Mehr als 90% der Welsarten sind solitäre, in 
Bodennähe lebende, nachtaktive, räuberische Süßwasserfische. Viele Callichthyidae, 
Pangasiidae, Schilbeidae und manche Siluridae sind tagaktiv, schwarmbildend und 
Kleintier- oder Pflanzenfresser. Die Loricariidae ernähren sich vorwiegend von 
Pflanzen und Detritus. Einige Ariidae, Aspredinidae und Plotosidae leben im Meer 
oder Brackwasser (Franke, 1985). Die Plotosidae zeigen ım Juvenilstadium ein 
besonders stark ausgeprägtes Schwarmverhalten (Burgess, 1989). Als Ostariophysi 
sensu stricto (Otophysi sensu Rosen & Greenwood, 1970) haben alle Welse einen 
Weberschen Apparat (Weber, 1820; Wright, 1884; Bridge & Haddon, 1889, 1893, 
1894; Sörensen, 1895; Chranilov, 1929; Alexander, 1964, 1965; Chardon, 1968). Er 
verbindet die Schwimmblase mit dem Innenohr und dient der Steigerung der 
Hörschärfe (von Frisch & Stetter, 1932). In vielen Welsfamilien gibt es Species, die 
in ihrer Epidermis Schreckstoffzellen führen (Pfeiffer, 1960, 1970, 1977). Beim 
Schreckstoff (von Frisch, 1938), der diesen Zellen entstammt (Pfeiffer, 1960; Smith, 
1973), handelt es sich um ein Pheromon (Karlson & Lüscher, 1959), das bei Frei- 
setzung durch Hautverletzung eine gesteigerte Aufmerksamkeit und eine Flucht- 
reaktion der Artgenossen auslöst. Dieses Verhalten wurde von Karl von Frisch (1938, 
1941a, b) an der Elritze (Phoxinus phoxinus, Cyprinidae) entdeckt und als Schreck- 
reaktion bezeichnet. Während die Schreckreaktion ein Merkmal aller Ostariophysi 
sensu lato, d.h. Gonorynchiformes (= Anotophysi, Rosen & Greenwood, 1970) plus 
Otophysi, ist (Pfeiffer, 1967), tritt der Webersche Apparat nur bei den Otophysi auf 
und ist somit das phylogenetisch jüngere Merkmal. Bei einigen Species oder Familien 
mit besonderer Lebensweise wurde die Fähigkeit zur Schreckreaktion sekundär 
teilweise oder vollständig rückgebildet (Pfeiffer, 1963a). So fehlt den Serrasalminae 
und Mylinae nur die Fähigkeit zur Schreckreaktion (im ZNS), obwohl sie in ihrer 
Epidermis Schreckstoffzellen (histologisch sichtbar) und damit Schreckstoff (im 
Verhaltensexperiment nachgewiesen) führen (Pfeiffer, 1962a, 1963a; Markl, 1968). 
Die Gymnotoidei und die Welsfamilien Aspredinidae und Loricariidae haben dagegen 
im Laufe ihrer Stammesgeschichte auch die Schreckstoffzellen und damit den 
Schreckstoff rückgebildet (Pfeiffer 1963a). Wie die meisten Knochenfische (Tele- 


168 ANDREAS HEYD & WOLFGANG PFEIFFER 


ostei) besitzen fast alle Welse (mit Ausnahme der Aspredinidae) Mauthner-Axone in 
ihrem ZNS und damit eine Mauthner-Reaktion, mit der auch ihre Schreckreaktion 
beginnt (Pfeiffer et al., 1986; Göhner & Pfeiffer, 1996). 

Etwas Besonderes innerhalb der Ostariophysi ist die Fähigkeit zahlreicher 
Welsarten, auf unterschiedliche Weise Laute zu erzeugen. Bei mindestens 8 Familien 
kommen Trommelmechanismen vor, und noch häufiger treten Stridulationsapparate 
auf. Innerhalb mehrerer Familien gibt es sogar beide Mechanismen nebeneinander. 
Ein Trommelmuskelapparat oder Springfederapparat ist bekannt von den Ariidae, 
Auchenipteridae, Bagridae, Doradidae, Malapteruridae, Mochokidae, Pangasiidae und 
Pimelodidae (Müller, 1842, 1857; Sörensen, 1884, 1895; Tavolga, 1962; Abu-Gideiri 
& Nasr, 1973; Kastberger, 1977, 1978; Kratochvil er al., 1980; Schachner & Schaller, 
1982; Ladich & Fine, 1994). Die Schwimmblase verstärkt als Resonanzraum die 
Vibration der mit ıhr direkt oder indirekt verbundenen Trommelmuskeln. 

Bei der Stridulation werden durch Gegeneinanderreiben von Knochenele- 
menten knarrende Geräusche erzeugt. Am weitesten verbreitet ist die Stridulation mit 
den Brustflossen. Sie ist nachgewiesen für die Arıidae, Aspredinidae, Auchenipter- 
idae, Bagridae, Callichthyidae, Doradidae, Heteropneustidae, Ictaluridae, Loricari- 
idae, Mochokidae, Pangasiidae und Pimelodidae. Die Laute entstehen durch Be- 
wegung des massiven Kopfes des ersten Brustflossenstrahls (Stachel), der einen 
Knochenkamm mit Querrillen und Höckern trägt und in eine gebogene Kerbe 
(Gelenkpfanne) im Cleithrum (Teil des Schultergürtels) eingepaßt ist. Wird dieser 
Knochenkamm gegen den Boden der Gelenkpfanne gepreßt und gleichzeitig bewegt, 
entstehen knarrende Geräusche (Geoffroy Saint-Hilaire, 1829; Dufossé, 1874; 
Haddon, 1881; Sörensen, 1884; Villwock, 1960; Winn, 1964; Pfeiffer & Eisenberg, 
1965; Agrawal & Sharma, 1965; Gainer, 1967; Abu-Gideiri & Nasr, 1973; Brousseau, 
1976; Kastberger, 1977, 1978; Kratochvil et al., 1980; Kratochvil & Völlenkle, 1981; 
Schachner & Schaller, 1982: Fine er al., 1996, 1997: Kaatz, 1999). Als Besonderheit 
ist von einigen Sisoridae die dorsale Stridulation (wie von Geoffroy Saint-Hilaire, 
1829 an Synodontis schall irrtümlich vermutet) bekannt. Hierbei reibt eine feilen- 
förmige Struktur an der Rückenflossenbasis durch Vor- und Rückwärtsbewegung 
gegen die gerippte Oberfläche der verwachsenen Neuratfortsätze des Wirbelkom- 
plexes (Haddon, 1881; Mahajan, 1963; Alexander, 1965). 

Bei allen Fischen mit besonderen lauterzeugenden Organen handelt es sich um 
Knochenfische (Osteichthyes), meist Teleostei. Bereits Aristoteles erwähnt sechs 
lauterzeugende Fischarten (Aristoteles, auch zitiert bei Cuvier & Valenciennes, 1840, 
Müller, 1857 und Sörensen, 1884, 1895). Es handelt es sich dabei um Species aus 
dem Mittelmeer. Geoffroy Saint-Hilaire (1829) beschrieb als erster die Lauterzeugung 
durch Stridulation eines Welses, nämlich Synodontis schall (Bloch & Schneider, 
1801) aus dem Nil (Daget er al., 1986). Seine richtige Aussage wurde von Cuvier & 
Valenciennes (1840) ohne eigene Beobachtung abgestritten, doch von Johannes 
Müller (1857) bestätigt. Erst knapp 100 Jahre nach Geoffroy Saint- Hilaire (1829) hat 
Karl von Frisch (1923) mit der von ihm in die Sinnesphysiologie und Verhaltens- 
forschung eingeführten genialen Dressurmethode das Hörvermögen für Fische nach- 
gewiesen, und zwar am nordamerikanischen Zwergwels Ameiurus nebulosus, "der 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 169 


kommt, wenn man ihm pfeift" (wofür der geblendete Fisch mit Futter belohnt wurde). 
Im Gegensatz zu den experimentellen Untersuchungen über Hörvermögen und 
Lauterzeugung gehen die ersten anatomischen Befunde und Beobachtungen an den 
lauterzeugenden Organen von Welsen weit mehr als 100 Jahre zurück (Müller, 1842; 
Agassiz, 1852; Reissner, 1859; Dufossé, 1874; Haddon, 1881; Wright, 1884; 
Sorensen, 1884, 1895, 1898; Bridge & Haddon, 1889, 1893, 1894). 

Eine Tierart bedarf, um zu überleben, des Schutzes ihrer Individuen vor dem 
Gefressenwerden durch Beutegreifer. Ein Schutz vor räuberischen Überfällen kann 
gegeben sein durch die Warnung der Artgenossen, beispielsweise durch ein Alarm- 
pheromon wie den Schreckstoff, und ist besonders bei wehrlosen, tagaktiven und 
schwarmbildenden Fischen zu erwarten. Im Gegensatz zu den meisten Species der 
Cyprinidae und Characiformes ist es gewöhnlich schwierig, die Schreckreaktion für 
die Siluroidei nachzuweisen (Pfeiffer, 1960, 1963a, 1967). Schutz (1956) erzielte an 
Ameiurus nebulosus acht und an Corydoras paleatus zwei positive Ergebnisse, ohne 
sie als Schreckreaktion zu deuten, trotz seines Hinweises, daß diese Welse als Jung- 
fische in Gruppen leben. Pfeiffer (1960) bestätigte die von Schutz (1956) an Ameiurus 
und Corydoras erhobenen positiven Befunde und wies die Schreckreaktion der Welse 
erstmalig an dem schwarmbildenden, tagaktiven Glaswels Kryptopterus bicirrhis 
(Siluridae) nach. Seitdem wurde die Schreckreaktion dieser und weiterer Species 
(Pangasius sutchi - Pangasiidae, Eutropiellus vandeweyeri - Schilbeidae und dem 
marinen Plotosus lineatus - Plotosidae) videotechnisch analysiert (Pfeiffer er al. 1986, 
Heyd & Pfeiffer, in Vorbereitung). Positive Ergebnisse liegen ferner für Angehörige 
der Mochokidae und Pimelodidae vor (Pfeiffer, 1963a). 

Eine völlig andere Möglichkeit des Schutzes ist die Abschreckung des An- 
greifers, beispielsweise akustisch durch Stridulationslaute oder Knarren. Sie ist 
hauptsächlich bei solitären, wehrhaften Species zu erwarten, wie den stacheltra- 
genden, meist nachtaktiven Welsen. Es war daher naheliegend zu prüfen, ob bei 
Welsen ein Zusammenhang zwischen Lauterzeugung und Schreckreaktion besteht, 
wie von Pfeiffer & Eisenberg (1965) vermutet. Trotz zahlreicher Sammelreferate über 
die Schreckreaktion einerseits (Pfeiffer, 1962b, 1963b, 1966, 1974, 1977, 1982: 
Smith, 1977, 1982, 1986, 1992; Solomon, 1977; Liley, 1982; Stabell, 1996) und die 
Lauterzeugung andererseits (Harden-Jones & Marshall, 1953; Tavolga, 1960, 1971a, 
1977; Schneider, 1961, 1967; Winn, 1964; Popper & Fay, 1973, 1993; Fine et al., 
1977; Hawkins & Myrberg, 1983; Bass, 1989; Schellart & Wubbels, 1998; Tyack, 
1998) wurde dieser Frage bisher keine Aufmerksamkeit geschenkt. Da sich die 
bisherigen Untersuchungen auf nur wenige Arten und Familien beschränken, haben 
wir weitere Species bezüglich ihrer Fähigkeit zur Lauterzeugung und deren Mecha- 
nismen studiert. 


MATERIAL UND METHODEN 


Bezüglich ihrer Lauterzeugung wurden 19 Species aus 8 Familien der Welse 
untersucht (Tabelle 1). Die Fische wurden als Jungtiere vom Großhandel bezogen; 
alle waren Wildfänge, ausgenommen Corydoras paleatus. Sie wurden in 200L- 


170 ANDREAS HEYD & WOLFGANG PFEIFFER 


Aquarien in Leitungswasser bei 24 - 26 °C Wassertemperatur gehältert (LD 12:12). 
Die Fütterung erfolgte je nach Species 1-2 mal täglich mit Flockenfutter, Futter- 
tabletten, roten Mückenlarven (Frostfutter) und Rinderherz. 


LAUTERZEUGUNG DURCH STRIDULATION: 


Die Versuche begannen nach 6 Wochen Quarantänezeit und fanden in einem 
ruhigen Raum statt. Die Welse wurden nach einer Eingewöhnungszeit von etwa einer 
Stunde aus einem kleinen Transportaquarium gefischt, an einer Brustflosse und am 
Körper gefaßt und in 5-10 cm Abstand vor ein Mikrophon (Uher M 517) gehalten. 
Die bei Bewegung der freien Brustflosse erzeugten Laute wurden mit einem 
Tonbandgerät (Uher report stereo) bei 19 cm/sec Bandgeschwindigkeit aufgezeichnet. 
Die einzelnen Fische wurden am selben Tag höchstens zweimal aus dem Aquarium 
genommen, wobei eine mindestens halbstündige Pause eingehalten wurde. Wenn ein 
Wels keine Laute erzeugte, wurde er später zu unterschiedlichen Tageszeiten, auch 
am Abend, wieder geprüft. 

Mittels eines Sonagraphen wurden von den Tonbandaufnahmen Sonagramme 
erstellt (Sonagraph Typ 6061, Papier Type B/65 Sonagram, Kay Elemetrics Corp., 
Pine Brook, NJ, USA), wobei die Registrierdauer 1,2 Sekunden im Frequenzbereich 
160-16000 Hz betrug. Durch Darstellung der Intensitätsverteilung konnte eine 
Frequenzanalyse der Laute durchgeführt werden. Die Tonbandaufnahmen wurden 
möglichst vollständig in Sonagramme übersetzt, aus denen die Längen der Laute und 
der Intervalle (Pausen zwischen den Lauten) bestimmt und in Millisekunden (ms) 
umgerechnet wurden. Die Mebwerte für jedes Individuum und für die Summe aller 
Angehörigen derselben Species wurden statistisch bearbeitet. Dabei wurde innerhalb 
derselben Art nicht zwischen Ab- und Adduktionslauten unterschieden. Ferner konnte 
durch Abspielen der Tonbänder mit reduzierter Geschwindigkeit die Anzahl der 
registrierten Laute bestimmt werden. Der Quotient aus Lautanzahl und zugehöriger 
Aufnahmedauer ergibt ein relatives Maß für die Aktivität (den Erregungsgrad) des 
Versuchsfisches. 


LAUTERZEUGUNG DURCH SCHWIMMBLASENMECHANISMEN: 


Zur Lautaufnahme wurden die folgenden Geräte eingesetzt: Panasonic NV-FS 
200 (SVHS-Videorecorder mit Long-Play-Aufnahmemôglichkeit); Videocassetten: 
Sony VHS E-240; Dyn. Mikrophon Vivanco- DM 22 (Frequenzbereich 60-12000 Hz, 
Impedanz 600 Ohm, Nierencharakteristik). Für die Unterwasseraufnahmen war dem 
Mikrophon ein Gefrierbeutel übergestülpt und wasserdicht abgeklebt. Dadurch wur- 
den die eingegebenen Frequenzen weder an Luft noch unter Wasser beeinflußt, wie 
der folgende Versuch gezeigt hat: mit einem Frequenzgeber (SINE/SQUAR Oscil- 
lator Li Interlab SQ 10 mit einem 50 Ohm/0,2 Watt Lautsprecher) wurden definierte 
Töne erzeugt und über das Mikrophon unverpackt oder verpackt registriert. Diese 
Aufnahmen wurden mit einem Frequenzzähler bzw. Frequenzanalyse-PC-Programm 
(SONA-PC, B. Waldmann, Zoologisches Institut der Universität Tübingen) analy- 
siert. Die Frequenz war stabil, das Mikrophon zeigte eine Richtcharakteristik. Mit 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 171 


zunehmender Entfernung vom Mikrophonkopf sank die Intensität der Töne, abhängig 
von der jeweiligen Frequenz. Die Versuche wurden unter den folgenden Bedingungen 
durchgeführt: Aquariengröße 80x50x50 cm, Wassertemperatur 27-28°C, Lichtphase 
7-19 Uhr (Leuchtstoffröhre); Fütterung Ix täglich am Vormittag. Die Fische wurden 
jeweils eine Woche eingewöhnt. 

Versuch A: Zur Bestimmung der Tag-Nacht-Lautaktivität wurde im Abstand 
von drei Wochen an zwei verschiedenen Gruppen aus je 4 Exemplaren von Arius 
seemani (Gesamtlänge 205-240 mm) die Lautaktivität drei Tage (72 Stunden) lang 
ohne Unterbrechung auf Videoband registriert. Der Aquarienraum wurde nur zum 
Programmieren des Recorders, Wechseln der Cassetten und zur Fiitterung betreten, 
um die Welse so selten wie möglich zu beunruhigen. Zur Bestimmung der Laut- 
aktivität wurden die Bandaufnahmen abgehört, wobei von jeder Stunde drei 5- 
Minuten-Abschnitte ausgewählt und jeweils die Laute gezählt wurden. Aus diesen an 
den drei Aufnahmetagen immer zur selben Stunde ermittelten Werten (n = 9) wurde 
der Median berechnet (= mittlere Lautaktivität in 5 Minuten). Dieser wurde mit 12 
multipliziert und dient als geschätztes Maß für die Lautaktivität während der 
betreffenden Stunde. 

Versuch B: Nach Abschluß der beiden Langzeitversuche wurden nachmittags 
drei Welse aus der zweiten Gruppe entnommen und die Laute des im Aquarium 
verbliebenen Individuums sofort nach dieser Störung bzw. nach einer Beruhigungs- 
phase von 15 Minuten registriert. Von diesen Aufnahmen wurden jeweils 20 Laute 
mit dem Sona-PC-Programm in bezug auf Frequenzanteile (Fourier-Analyse) sowie 
Laut- und Intervalldauer ausgewertet. 

Für die vergleichend-morphologischen Untersuchungen dienten je 3 Arius 
seemani (215-260 mm Gesamtlänge), Pimelodus pictus (105-110 mm) und Pimelo- 
della gracilis (110-125 mm). Von jeder Species wurden nach Tötung mit MS 222 
(Sigma Chemical Co.) ein Exemplar frisch präpariert und zwei in Formalin (1:10 
verdünnt) fixiert. Alle Fische wurden ventral geöffnet und geprüft auf den Besitz 
eines Fensters in der Muskulatur jeder Körperseite (“the lateral cutaneous areas”: 
Bridge & Haddon, 1893; Alexander, 1964, 1965, 1966). Ferner wurden Lage, Größe 
und Form ihrer Schwimmblase notiert. Schließlich wurde nach einem Trommel- 
muskelapparat gesucht. Während der Präparation wurde mit einer ZEISS-Auf- 
setzkamera M 35 am ZEISS-Standardbinokular fotographiert (Filmmaterial: Agfa 
HDC 100/21, Fujichrome 50/18, Kodak 100/21). 


ERGEBNISSE 
STRIDULATION 

Alle Species reagierten auf Berührung mit Zappeln, Schwanzschlagen, Auf- 
richten der Rückenflosse sowie mit Abspreizen und Arretieren oder Bewegungen der 
Brustflossen. Bei letzterem erzeugten 13 der 18 untersuchten Species Stridulations- 
laute (Tab. 1, 2). Am lautesten waren die beiden Synodontis-Species, Agamyxis und 
Glyptoperichthys, während Peckoltia besonders leise stridulierte. Die Angehörigen 
derselben Art waren unterschiedlich erregt: die einen begannen schon beim Heraus- 
fischen zu stridulieren, die anderen erst beim Anfassen, einige blieben auch bei 


172 ANDREAS HEYD & WOLFGANG PFEIFFER 


TABELLE | 


Familienzugehörigkeit der 19 untersuchten Welsarten, in Tab. 2 und Abb. 6-9 verwendete 
Abkürzungen (Abk.), Anzahl der auf Stridulation geprüften Exemplare (N) und deren Gesamt- 


länge. 

* es könnte sich um Peckoltia vittata (Steindachner, 1882) handeln. 
** Syn. Pimelodella meesi. *** nur auf Trommelmechanismen untersucht. 
Familie / Species Abk. N Länge (mm) 
Ariidae 
Arius seemani Günther, 1864 As 12 60-85 
Aspredinidae 
Dysichthys coracoideus Cope, 1874 De 5 80-100 
Bagridae 
Hemibagrus nemurus (Valenciennes, 1839) 4 110-122 
Mystus gulio (Hamilton, 1822) Mg 4 63-67 
Mystus vittatus (Bloch, 1794) Mv 3 55-62 
Callichthyidae 
Corydoras paleatus (Jenyns, 1842) 13 40-50 
Dianema urostriata De Miranda-Ribeiro, 1912 Du 5 40-55 
Doradidae 
Agamyxis flavopictus (Steindachner, 1908) Af 3 53-65 
Loricariidae 
Ancistrus cf. dolichopterus Kner, 1854 3 55-70 
Glyptoperichthys gibbiceps (Kner, 1854) Gg 3 100-120 
Panaque nigrolineatus (Peters, 1877) 2 55-60 
Parancistrus aurantiacus (Castelnau, 1855) | 80 
Peckoltia pulcher (Steindachner, 1915) * Pp 3 73-90 
Pseudacanthicus spinosus (Castelnau, 1855) 3 70-95 
Mochokidae 
Synodontis ocellifer Boulenger, 1900 So 3 105-130 
Synodontis schoutedeni David, 1936 Ss 3 80-100 
Pimelodidae 
Brachyrhamdia meesi Sands & Black, 1985** Bm | 70 
Pimelodella gracilis *** (Cuvier & 

Valenciennes, 1840) = = 

Pimelodus pictus Steindachner, 1876 4 75-90 


wiederholtem Fangen ruhig. Eine hohe Stridulationsaktivität zeigten die beiden 
Mystus- und Synodontis-Arten sowie Agamyxis flavopictus und Peckoltia pulcher 
(Tab. 2). Der Erregungsgrad aller untersuchten Individuen lag im Durchschnitt 
zwischen 0,9 und 2,5 Lauten pro Sekunde. Nur bei 5 Species stridulierten alle 
Individuen; von 66 stridulierfähigen Individuen aller Species blieben 24 (36%) auch 
bei wiederholten Störungen stumm (Tab. 2). Stridulation im Wasser trat nur selten 
auf. Einmal lärmte ein Agamyxis im Aquarium, als er mit zwei Artgenossen in die 
Quere kam. Ein Pimelodus stridulierte im Becherglas, mit dem er aus dem Aquarium 
genommen worden war. Dysichthys beantwortete das Anfassen im Wasser mit 
Körpervibrationen. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 173 


TABELLE 2 


Ergebnisse der Untersuchungen zur Stridulation (Abkürzungen der Speciesnamen vgl. Tabelle 
1). Nur stridulierende Species sind hier aufgelistet. 

S: Stridulationsweise (Ab = Abduktion, Ad = Adduktion). 

n (N): Anzahl n der stridulierenden Individuen von allen geprüften Exemplaren (N) einer 
Species; E: Erregungsgrad (Anzahl Laute / sec., Mittelwerte + Standardabweichung); L: Laut- 
dauer (Medianwerte) in ms; P: Intervalldauer (Medianwerte) in ms; F: Frequenzschwerpunkte 
der Laute in KHz. 


L P F 


Sea SÌ n(N) E 

As Ab 6 (12) IR 0.5 30.2 77.4 1-3; 4-5; 10-11 
Bm Ab 170) 0.9 52.8 TS 2-4; 6-8 
Du Ab SO) 1.0+0.6 41.5 Wal 0.5-2; 3-4 
Gg Ab 313) M 087] SS 2302 1.5-2.5; 4-5 
Pp Ab 16) 1.6 + 0.6 67.9 130.2 4-6; 9-12 
Af Ab+Ad 3:6) 1.4 +0.4 101.9 56.6 2-3; 5-6 
De Ab+Ad 4 (5) 19122086 83.95 41.5 0.5-2 
Mg Ab+Ad 3 (4) 14+1.2 45.3 34 3-4; 7-8 
Mv Ab+Ad 348) 1.4 +0.4 34.9 34 3-5; 8-10 
So Ab+Ad 28) Ses 10 61.35 Sell 1-2; 3.5-4.5 
Ss Ab+Ad 2 (3) 29-13 49.1 23893 0.5-2 


Die Stridulationslaute entstanden je nach Species auf unterschiedliche Weise. 
Während Arius, Brachyrhamdia, Dianema, Glyptoperichthys und Peckoltia nur bei 
Abduktion (Abspreizen) der Brustflosse Laute erzeugten, geschah dies bei Agamyxis, 
Dysichthys, den Mystus- und den Synodontis-Arten bei Abduktion und Adduktion 
(Heranziehen) der Brustflosse (Tab. 2). Lauterzeugung nur bei Adduktion wurde nie 
beobachtet. Die Geräusche bestehen aus Serien einzelner Laute, welche wiederum aus 
Einzelknacks aufgebaut sind, die bei kürzeren Lauten dichter aufeinander folgen als 
bei längeren. Diese Einzelknacks sind bei Arten mit Abduktionslauten ziemlich 
gleichmäßig angeordnet und die Laute einer Serie ähnlich gestaltet (Abb. 1, 2). Teil- 
weise können anhand der Sonagramme die Abduktionslaute von den Adduktions- 
lauten an ihrer unterschiedlichen Dauer bzw. der Dichte der Einzelknacks unter- 
schieden werden (Abb. 3, 4, 5). Die Laute zeigen keinen harmonischen Aufbau, 
sondern eine breite Frequenzverteilung. Die Bereiche größter Energie sind in den 
Sonagrammen an den stark geschwärzten Stellen (Banden) erkennbar (Abb. 1-5). Oft 
treten zwei oder drei Banden auf, wobei die niedrigste Frequenz den Hauptbereich 
darstellt. Die Hauptfrequenzanteile liegen bei 500 - 6000 Hz (Tab. 2). 

Aufgrund individueller Unterschiede (Abb. 6) ergeben sich in der Zusammen- 
fassung für jede Species teilweise große Schwankungen der Laut- und noch mehr der 
Pausenlängen (Abb. 7). Kurze Laute im Bereich bis 50 ms erzeugen Arius, Dianema, 
Mystus gulio, M. vittatus sowie Synodontis schoutedeni. Bei den restlichen Species 
betragen die mittleren Lautlängen 50 - 100 ms (Tab. 2). Die größte Variabilität der 
Lautlänge zeigen Agamyxis, Dysichthys, Glyptoperichthys und Peckoltia, wobei ein- 
zelne Laute mehr als 150 ms dauern. Auch das einzige Exemplar von Brachyrhamdia 


174 


ANDREAS HEYD & WOLFGANG PFEIFFER 


Mm 


2 4 
az Be 

= 2 

È è iu À 

® il 

Es 3 | 

<= & | 


sie va ve 
IT] 


eo 
rn re 


ur 
| 


| [ll | 
STILI 


di i i | im citi il 
È > ji LA Mi dl) A 
2 2. I Ht 1 Mi 


100 ms 


ie 


1 Be 


ABB. | 


Ausschnitt je eines Sonagramms von: a) Arius seemani, b) Brachyrhamdia meesi, c) Dianema 
urostriata. Abszisse: Zeitskala in ms. Ordinate: Frequenzskala 0-16 kHz in 1-kHz-Stufen. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 175 


| N 
| | 
ti); 


| 


ABB. 2 


Ausschnitt je eines Sonagramms von: a) Glyptoperichthys gibbiceps, b) Peckoltia pulcher. 
Achsen wie in Abb. 1. 


weist bei kürzerer Lautdauer eine große Streuung auf (Abb. 7a). Die Pausenlänge ist 
am geringsten und streut am wenigsten bei den zwei Synodontis-Arten und Mystus 
vittatus; sie ist am größten bei Glyptoperichthys und streut am breitesten bei Dianema 
und Dysichthys (Abb. 7b). 

Während ein Zusammenhang zwischen Erregungsgrad und Laut- bzw. 
Pausenlängen nicht ersichtlich ist (Tab. 2), zeigt sich im Verhältnis von Lautlänge zu 
Pausenlänge eine Abhängigkeit von der Art der Lauterzeugung (Abb. 8). Bei Species, 
die Ab- und Adduktionslaute erzeugen, sind die Laute mindestens ebenso lang wie die 
Pausen (Verhältnis meist größer als 1); bei Species mit Abduktionslauten übersteigt 
die Intervalldauer die Lautdauer teilweise weit (Verhältnis kleiner als 1). 


176 


ANDREAS HEYD & WOLFGANG PFEIFFER 


my 


ni 


N | 


[LI 


ME 


wi 


m 


% 


Hebe ly 


| 


8 
ie 


Ae 


ta 


LI | 
È Wa 


si He 


ee HIN ern i 


IM | || “| di x È a 


ds 
| 


i 


I 
Wr 


ee 


| fai 


HR ©; 


100 ms 


ABB. 3 


Ausschnitt je eines Sonagramms von: a) Mystus gulio, b) Mystus vittatus. Achsen wie in Abb. 1. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 11/7 


Hi 
N Il 


& È DT | ev h 
trad à 
L'ART 4 vr. ry 
n SR = dle, m nm oe 
L- = a a Me 
= = = eee == = = 
a Meet Pr. Zi ee SII 


ic 


ti 16! 


kr 
an Tr 9 


: yi MN 
a | 
ae | ll ih, 
: i IN) | | i oil 
ri SANTI ne 


100 ms 


ABB. 4 


Ausschnitt je eines Sonagramms von: a) Synodontis ocellifer, b) Synodontis schoutedeni. 
Achsen wie in Abb. 1. 


Der Panzerwels Corydoras paleatus erzeugte Stridulationslaute auf folgende 
Art und Weise: wurde der Hinterkörper beidseitig gefaßt, warf der Fisch seinen 
Vorderkörper schnell hin und her. Dabei waren Laute hörbar, deren Sonagramme 
Stridulationslauten ähneln (Abb.9). Wurde der Fisch am Rumpf und an einer Brust- 
flosse festgehalten, blieb die andere Pectoralis arretiert. Im Gegensatz zur stridu- 
lierenden Peckoltia zeigten die Loricariidae Ancistrus cf. dolichopterus, Panaque 
nigrolineatus, Parancistrus aurantiacus und Pseudacanthicus spinosus aus derselben 
Unterfamilie Ancistrinae zu keiner Tageszeit Stridulation. Alle Individuen dieser 


178 ANDREAS HEYD & WOLFGANG PFEIFFER 


Mi (1 , , sn e Ms fe tue 
+ Aad igs È = 4 In 
3 i Hal 88, Brit DIR br O A ri 


ee eee 


b è € i 


1° | ih hs di L Fi; i | 
" I a i 


100 ms 


Lu 
| BA Bi 


BR 


| 4 | oti 
Rie? 1 Duo 


ABB. 5 


Ausschnitt je eines Sonagramms von: a) Agamyxis flavopictus, b) Dysichthys coracoideus. 


._ Achsen wie in Abb. I. 


Species, ausgenommen Peckoltia, verhielten sich beim Herausnehmen aus dem 
Wasser ähnlich. Sie spreizten und arretierten beide Brustflossen, richteten ihre 
Rückentlosse auf und führten Schwanzschläge aus. Im Gegensatz zu Pseudacanthicus 
klappten Ancistrus und Parancistrus zusätzlich ihre mit Hakenstacheln besetzten 
Interopercula nach außen. Panaque besitzt keine solchen Interopercula. Anders als die 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 179 


ms 
200 


600 


20 14 


300 31 


100 774010 19 32 2 44 


ABB. 6 


Dauer der Stridulationslaute (a) und der Pausen zwischen den Lauten (b) von 33 Individuen aus 
11 Species. Dargestellt sind Mittelwert, Standardabweichung, Minimum und Maximum sowie 
Anzahl der Laute (Ziffer). Abszisse: Species (Abkürzungen siehe Tab. 2; Pg = Pterygoplichthys 
gibbiceps = Synonym von Glyptoperichthys gibbiceps = Gg). Ordinate: Dauer in ms. Beachte 
die stärkere Dehnung der Ordinate in (a) gegenüber (b). 


180 ANDREAS HEYD & WOLFGANG PFEIFFER 


200 
150 


100 


600 


= 
200 Za 
BEER 


As Bm Du Gg Pp Af Dc Mg Mv So Ss 


ABB. 7 
Box-whisker-plots der Lautdauer (a) und der Pausendauer (b) der 11 Species. Abszisse: Species 
in Reihenfolge nach der Stridulationsweise: As-Pp Abduktion; Af-Ss Ab- und Adduktion. Ab- 
kürzungen siehe Tab. 2. Ordinate: Dauer in ms. Box: Median, 25. und 75. Perzentil. Whisker: 
10. und 90. Perzentil. Punkte: 5. und 95. Perzentil. 
beiden Bagridae der Gattung Mystus stridulierten vier wiederholt gepriifte Exemplare 
von Hemibagrus nemurus nicht, zeigten jedoch ein anderes bemerkenswertes Ver- 
halten. Sie erzeugten mehrmals hintereinander durch Auspressen von Luft aus ihren 
Kiemenspalten Quietschlaute von 26 - 56 ms Dauer, mit einer Hauptfrequenz von 1,5 
kHz (Abb. 10). Bei Pimelodus pictus waren wenige Stridulationslaute gleichzeitig mit 
Trommelmuskellauten zu hören. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 181 


EE 


As Bm Du Gg Pp Af Dc Mg Mv So Ss 


ABB. 8 


Verhaltnis von Lautdauer zu Pausendauer fiir jede Species, dargestellt als box-whisker-plots. 
Erläuterung siehe Abb. 8. 


TROMMELMECHANISMEN: 


Pimelodus pictus: alle vier Individuen erzeugten Laute, die an das Summen 
einer Fliege oder Hummel erinnern. Ein Exemplar stridulierte kurz außerhalb des 
Wassers, wobei gleichzeitig das Summen ertönte. Auch ein im Becherglas stridu- 
lierendes Exemplar summte. Die Summtöne waren etwa | sec lang und wurden mehr- 
mals hintereinander erzeugt. 

Arius seemani: beim Transport dieser Welse waren ebenso wie im Aquarium 
Laute vernehmbar, die sich wie „Popps“ anhörten und entfernt an das Gackern von 
Hühnern erinnerten. Auf Klopfen an das Aquarium stellten die Fische diese 
Geräusche für einige Sekunden ein. Manche Individuen ließen diese Laute auch 
außerhalb des Wassers hören, während sie gleichzeitig stridulierten. Diese Laute 
waren leiser als das durch Stridulieren erzeugte Knarren. 


ANDREAS HEYD & WOLFGANG PFEIFFER 


182 


[UT 


RR iii 


nee B 


mL 


BEA an fr 


ARBRE tt 18: 


100 ms 


ABB. 9 
on Corydoras paleatus (2 Exemplare). Achsen wie in Abb. 1. 


le 


Ausschnitte aus Sonaerammen v 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 183 


Li! 


1 
Fin 


ye 


Li 


| | 
À 
{ 


hi 
1 


# E 

= 
= a 
=. = 
= at ae È 
FA “= wu 
a & 


100 ms 


ABB. 10 


Zwei “Quietschlaute” eines Exemplars von Hemibagrus nemurus. a: Sonagramm. b: Frequenz- 
analyse des Sonagraphen; vertikal Frequenz, horizontal Amplitude. 


Versuch A: Die 4 Fische der ersten Gruppe begannen gleich nach dem 
Einsetzen in das Versuchsaquarium Trommellaute zu erzeugen, die in | m Entfernung 
vom Aquarium zu hören waren. Tagsüber lagen diese Welse entweder eng bei- 
sammen ruhig am Boden oder ruhten mit dem Kopf nach oben fast senkrecht an einer 
Aquarienwand. Bei Fütterung schwammen sie umher, danach ruhten sie wieder 
beieinander. Auch die zweite Gruppe gab sofort nach dem Einsetzen ins Versuchs- 
aquarium Trommellaute von sich. Sie verhielt sich insgesamt ruhiger als die erste 


184 ANDREAS HEYD & WOLFGANG PFEIFFER 


ABB. 11 


24-Stunden-Lautaktivität der beiden Gruppen (Gr. 1, Gr. 2) von Arius seemani. Dargestellt sind 
Medianwerte über 3 Tage. Abszisse: Uhrzeit. Ordinate: Anzahl der Laute. 


Gruppe und erzeugte bei Fütterung und beim Hantieren am Aquarium weniger Laute. 
Das Verhalten am Tag ähnelte dem der ersten Gruppe. Beide Gruppen zeigten in ihrer 
Lautaktivität einen ausgeprägten Tag-Nacht-Rhythmus (Abb. 11). Die Lautproduktion 
war am Tag deutlich höher als nachts und etwa | Stunde nach Einsetzen der 
Lichtperiode am höchsten. Die Lautaktivität nahm dann ab, um gegen 12 Uhr ein 
erneutes, aber kleineres Maximum zu erreichen. Am Nachmittag war ein weiteres, 
schrittweises Abklingen der Lautanzahl zu verzeichnen. Zwischen 19 und 7 Uhr 
(Dunkelphase) erzeugten die Welse nur wenige Laute. Bereits zwei Stunden vor der 
Lichtperiode (7 Uhr) intensivierten beide Gruppen ihre Lautaktivität. Die Lautserien 
waren am Morgen gehäuft, am Mittag vereinzelt und wurden oft nur von einem ein- 
zigen Fisch erzeugt. Die nachts registrierten Laute waren dagegen meist Einzel- 
geräusche, nur ausnahmsweise Serien. Bei Gruppe 1 war die Gesamtzahl der Laute in 
der Hellphase um etwa 50 %, nachts um etwa 23 % höher als bei Gruppe 2. 

Versuch B: Alle Laute des isolierten Exemplars zeigten gleiche Frequenz- 
spektren. Die Hauptfrequenzen lagen unter 500 Hz. Die maximale Intensität war bei 
140 - 200 Hz, hohe Intensitäten sind bei 250 - 400 Hz zu erkennen (Abb. 12). Die 
mittlere Lautdauer betrug 209,5 + 17,4 ms, die Intervalldauer 0,3 - 6 sec. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 185 


0 200 1000 ms 
ABB. 12 


Farbspektrogramm der Trommellaute von Arius seemani. Abszisse: Zeitachse in ms. Ordinate: 
Frequenz in kHz. Farbskala rechts: Intensitätsverteilung des Schalldruckpegels in dB. 


Struktur der Trommelmuskelapparate (Schwimmblasenmechanismen): 


Arius seemani besitzt im Bereich der vorderen Schwimmblasenkammer, ein 
laterales Fenster in der Muskulatur (“lateral cutaneous area”). Die Versteifung des 
Schädel-Hals-Bereiches wird durch den mit dem Basioccipitale fest verwachsenen 
Wirbelkomplex verstärkt. Eine knöcherne Lamelle des Epioticums ist mit dem 
Transversalfortsatz des 4. Wirbels fest verbunden. Der Wirbelkomplex wird von den 
Wirbeln 2 bis 7 gebildet. Die Wirbel 4 bis 7 formen mit ihren Transversalfortsätzen 
eine der Schwimmblase aufliegende Platte. Während die Transversalfortsätze 6 und 7 
nur starres Bindegewebe miteinander verbindet, sind die Fortsätze 4 und 5 
miteinander verwachsen. Der anteriore Teil des Transversalfortsatzes des 4. Wirbels 
ist zu einem langen, ventrad gebogenen Ast ausgezogen. Tavolga (1962) hat ihn zu 
Ehren seines Entdeckers Johannes Müller (1842) ,,Miiller’ sche Ramus“ genannt 
(Abb. 13). Dieser lanzenförmige Ramus liegt der Schwimmblase craniad auf und ist 
mit ihr über Bindegewebe verwachsen, das ihn bis zum Ansatz des Trommelmuskels 
überzieht. Dieser indirekte Trommelmechanismus wurde von Müller (1842) als 
„Springfederapparat“ (,,elastic spring apparatus“) bezeichnet. Er wird vom Trans- 
versalfortsatz des 4. Wirbels, dem Müller’schen Ramus und dem Trommelmuskel 
(Protractor) gebildet. Der Protractor hat seinen Ursprung am Posttemporale, dem 
Epioticum und dessen knöcherner Lamelle. Die Ansatzstellen des Muskels liegen am 
Vorderrand des Transversalfortsatzes des 4. Wirbels bis zu dessen Verbindung mit 


186 ANDREAS HEYD & WOLFGANG PFEIFFER 


ABB. 13 


Springfederapparat von Arius seemani (215 mm) der rechten Körperseite von lateral. P, 
Protractor mit blutrotem anterioren Anteil. El, knöcherne Lamelle des Epioticums. TP,, 
Transversalfortsatz des 4. Wirbels. MR, Müller’sche Ramus. 8:1. 


der Epioticum-Lamelle. Ein weiterer Teil des Muskels zieht zum proximalen 
Müller’schen Ramus. Der vordere Bereich des fächerförmigen Trommelmuskels 
erscheint am frischen Präparat stärker blutrot als der zum Transversalfortsatz des 4. 
Wirbels ziehende Ast. Beide Muskelbereiche lassen sich kaum voneinander trennen, 
da sie gemeinsam den Protractor bilden (Abb. 13). Die herzförmige Schwimmblase 
eines 260 mm großen Arius ıst 33 mm lang und erstreckt sich bis zum 8. Wirbel, 
eingerahmt vom Müller’schen Ramus, dem Wirbelkomplex und dessen Transver- 
salfortsätzen. Sie wird durch ein unvollständiges Diaphragma in eine anteriore und 
eine posteriore Kammer geteilt. Letztere wird durch zwei Transversalsepten weiter 
unterteilt. Die Ansatzstellen der Septen sind an der dorsalen und ventralen Schwimm- 
blasenwand verbreitert (Abb. 14). Der kleine Ductus pneumaticus zieht vom caudalen 
Ende der vorderen Schwimmblasenkammer zum Vorderdarm. 

Pimelodus pictus: Die vordere Schwimmblasenkammer grenzt teilweise an die 
Haut. Die Wirbelkörper 2 bis 5 sind miteinander verwachsen. Die Transver- 
salfortsätze der Wirbel 4 und 5 bilden zusammen eine starre, der Schwimmblase auf- 
liegende, leicht geschwungene Platte. Der Transversalfortsatz des 4. Wirbels teilt sich 
distal in einen anterioren und einen posterioren Ast, an dessen wulstartig verstärktem 
cranialem Rand der Trommelmuskel entspringt. Dieser Fortsatz ist gelenkig mit dem 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 187 


ABB. 14 


Schwimmblase von Arius seemani (260 mm, männlich), linke Körperseite, lateral geöffnet. a) 
anteriore Kammer, durch das Diaphragma (D) von der posterioren Kammer teilweise getrennt. 
8:1. b) posteriore Kammer. Deutlich sichtbar die Transversalsepten (TS) und das Diaphragma 
(D), das sich wie die TS and der dorsalen und ventralen Schwimmblasenwand verbreitert. 8:1. 


188 ANDREAS HEYD & WOLFGANG PFEIFFER 


ABB. 15 


Schwimmblasen von ventral. a) Pimelodus pictus (105 mm). Der blutrote Trommelmuskel 
(TM) überdeckt mehr als die Hälfte der Schwimmblase (SB). 8:1. b) Pimelodella gracilis 
(123 mm). Schwimmblase frei, kein Trommelmuskel. 8:1. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 189 


ABB. 16. Pimelodus pictus (Kopf nach links gelegen) a) Schwimmblase eines 105 mm langen 
Exemplars von links lateral geöffnet. Die anteriore großlumige Kammer ist von der posterioren 
Kammer durch ein Diaphragma (D) teilweise getrennt. Der hintere Schwimmblasenteil ist von 
Transversalsepten (TS) weiter unterteilt. 12:1. b) Schwimmblase eines 120 mm langen for- 
malin-fixierten Exemplars, von dorsal geöffnet. Deutlich sind die zur dorsalen Schwimm- 
blasenwand ziehenden Septen, die sich an ihren Ansatzstellen verbreitern. 8:1. 


190 ANDREAS HEYD & WOLFGANG PFEIFFER 


Posttemporale verbunden. Der paarige, durch große Blutgefäße versorgte und daher 
intensiv rote Trommelmuskel (Protractor) hat seinen Ursprung am Transversalfortsatz 
des 4. Wirbels und reicht über die Hälfte der ventralen Schwimmblasenwand. Er 
bedeckt die vordere Schwimmblasenkammer mit Ausnahme der “lateral cutaneous 
areas” und setzt am cranialen Ende der hinteren Kammer an. An seiner Ansatzstelle 
läuft der Trommelmuskel flach aus. Seine Ansatzareale sind S-förmig gekrümmt. Die 
herzförmige Schwimmblase erstreckt sich vom 2. bis zum 6. Wirbel und ist bei einem 
105 mm großen Exemplar 15 mm lang (Abb. 15). Sie wird durch ein unvollständiges 
Diaphragma in eine anteriore und eine posteriore Kammer unterteilt. Weil das 
Diaphragma von lateral-dorsal gelegenen Öffnungen durchbrochen wird, sind die 
beiden Kammern miteinander verbunden. Die posteriore Kammer wird durch Trans- 
versalsepten nur unvollständig weiter unterteilt. Die Septenbildung beginnt an der 
Ansatzstelle des Trommelmuskels und reicht bis zum caudalen Schwimmblasenende. 
Die Septen ziehen zur dorsalen und ventralen Schwimmblasenwand (Abb. 16). Der 
Ductus pneumaticus entspringt in der Mitte des caudalen Endes der vorderen 
Schwimmblasenkammer. 

Pimelodella gracilis besitzt ebenso wie Pimelodus pictus und Arius seemani 
in der lateralen Muskulatur beidseitig je ein ein Fenster, so daß ein Teil der vorderen 
Schwimmblasenkammer, nur von Haut überdeckt, in engem Kontakt mit dem 
Außenmedium Wasser steht. Die Wirbel 3 bis 5 sind miteinander verwachsen. Die 
Transversalfortsätze der Wirbel 4 und 5 bilden zusammen eine der Schwimmblase 
aufliegende Platte. Der anteriore Teil des Transversalfortsatzes des 4. Wirbels formt 
an seinem cranialen Ende einen Wulst, ähnlich wie bei Pimelodus. Ein Trommel- 
muskelapparat wurde bei Pimelodella jedoch nicht gefunden. Die herzförmige 
Schwimmblase erstreckt sich vom 2. bis zum 6. Wirbel und ist bei einem 123 mm 
großen Exemplar 8,5 mm lang. Sie wird von einem unvollständigen Diaphragma in 
eine anteriore und eine posteriore Kammer unterteilt. Das Diaphragma besitzt rechts 
und links je eine Öffnung, durch welche die beiden Kammern kommunizieren (Abb. 
17). Die posteriore Kammer ist mindestens doppelt so lang wie die anteriore und wird 
durch ein vertikales Längsseptum vollständig zweigeteilt. Weitere Transversalsepten 
wurden, im Gegensatz zu Pimelodus, nicht gefunden. Der Ductus pneumaticus 
entspringt an der Schnittstelle des Längsseptums mit dem Diaphragma. 


LAUTERZEUGUNG UND SCHRECKREAKTION 


In Tabelle 3 sind diejenigen Welsfamilien aufgelistet, die auf den Besitz von 
Schreckstoffzellen und Schreckreaktion bzw. ihre Fähigkeit zur Lauterzeugung unter- 
sucht worden sind. Ferner zeigt diese Übersicht das Vorkommen sowie die Anzahl 
von Gattungen und Species nach Nelson (1994) und Teugels (1996). Der Reihenfolge 
der Familien liegt ein Cladogramm von de Pinna (1998) zugrunde, wobei die 
„niedrigen“ Taxa zuerst genannt sind. Während 20 der 23 dargestellten Taxa Schreck- 
stoffzellen besitzen, ist die Schreckreaktion nur bei 5 Welsfamilien durch Video- 
aufnahmen dokumentiert und bei 4 weiteren beobachtet worden. Stridulationsmecha- 
nismen treten bei 14, Trommelmechanismen bei 8 Familien auf. Beide Lauter- 
zeugungsmechanismen gemeinsam kommen bei 7 der „höheren“ Taxa vor. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 19] 


ABB. 17. Pimelodella gracilis (Kopf nach rechts gelegen). a) Schwimmblase eines 123 mm 
langen Exemplars, rechte Körperseite lateral geöffnet. Vordere und hintere Kammer sind durch 
ein Diaphragma (D) teilweise getrennt. Die hintere Kammer wird von longitudinalem Septum 
(LS) vollständig in zwei Hälften unterteilt. 12:1. b) Schwimmblase eines 100 mm Exemplars, 
ventral geöffnet. Die seitlichen Öffnungen im Diaphragma sind im Vergleich zu Pimelodus 
pictus großlumiger, weitere transversale Septen fehlen. 8:1. 


192 ANDREAS HEYD & WOLFGANG PFEIFFER 


TABELLE 3 


Vorkommen, Anzahl der Gattungen (G.) und Species (Spec.) nach Nelson (1994) und Teugels 
(1996) sowie das Auftreten von Schreckstoffzellen (SZ), Schreckreaktion (SR), Stridulation 
(ST) und Trommelmechanismen (TM) bei verschiedenen Familien und Unterfamilien der 
Welse. Reihenfolge der Familien nach einem Cladogramm in de Pinna 1998, p. 289; 
Verwandtschaftsgruppen mit @ markiert. * : Unterfamilie der Pimelodidae = Rhamdiinae in 
Teugels 1996. **: Familie bei de Pinna 1998 nicht aufgelistet. 

Abkürzungen: Ab = Abduktion; Ad = Adduktion; AF = Afrika; AS = Asien; D = Rückenflosse; 
E = Europa; MA = Mittelamerika; MAR = marin; NA = Nordamerika; PA = Pazifik; SA = 
Südamerika; SW = Süßwasser; TS = tropisch-subtropisch; V = Videoaufnahmen von SR vor- 
handen; + = vorhanden; (+) = SR gesehen; + ? = vorhanden, Mechanismus nicht beschrieben; ? 
= fraglich; -- = nicht vorhanden. 


G./ Spec. 


G./Spec. SZ SR ST ™ 


Familie Vor- 


kommen Nelson 1994 Teugels 1996 
@ Cetopsidae SW/SA 4/12 4/14 + 
@ Loricariidae SW /SA, MA 80 / > 550 88/651 -- -- Ab 
Callichthyidae SW /SA, MA 71130 7/144 + V Ab 
Trichomycteridae SW / MA, SA 36 / 155 40/154 + 
Amphiliidae SW / AF 7/47 9/60 + 
Sisoridae SW/AS 20/85 20/98 + D 
@ Aspredinidae SW/SA 10 / 32 10 / 32 -- -- Ab+Ad 
© Heptapterinae* SW/SA : 18 / 220 Ab 
@ Ictaluridae SW/NA 7145 7145 + (+) Ab/Ab+Ad 
@ Doradidae SW/SA 35 / 90 37 / 94 + Ab+Ad + 
Auchenipteridae SW / SA 21/60 20/61 + + + 
Mochokidae SW / AF 10 / 167 11/177 + (+) Ab+Ad + 
Ariidae MAR (SW) / TS 14 / 120 14/148 + Ab + 
@ Siluridae SW/E, AS 12/100 9/60 + V 
Malapteruridae SW / AF 12 10/73 + + 
Chacidae SW / AS 193 13 + 
Plotosidae MAR, SW / AS, PA 9/32 932 + V ? 
Clariidae SW / AF, AS 13 / 100 15 / 92 Et (Gr) ? 
@ Pangasiidae SW / AS 2/21 2/2] + V + + 
Schilbeidae SW / AF, AS 18/45 13751 + V 
@ Bagridae SW / AF, AS 30/210 [Sys SEE Ab+Ad + 
@ Pimelodinae SW / MA, SA 56 / 300 931/323 ab (©) Ab + 
@ Heteropneustidae** SW / AS 17/22 YE + apa 
DISKUSSION 
LAUTERZEUGUNG: 


Ariidae: Die Unterwasserlaute von Arius felis (Synonym Galeichthys felis) stu- 
dierte Dobrin (1947). Diese Species erzeugt sowohl Laute mit dem Springfeder- 
apparat (Burkenroad, 1931) als auch Stridulationsgeräusche. Frisch gefangene A. felis 
brachten bei Bewegung der Brustflossen 30 - 50 ms lange Laute im Frequenzbereich 
2 - 4 kHz hervor (Tavolga, 1960). Arius seemani produziert bei Abduktion der 
Brustflossen Stridulationslaute, deren Dauer und Frequenzanteile ebenfalls in diesem 
Bereich liegen. Die Lauterzeugung mittels Springfederapparat wurde von Tavolga 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 193 


(1971b, 1976, 1977) eingehend untersucht. Arius felis erzeugt fast dauernd Trom- 
mellaute der Frequenz 100 Hz und benützt sie als primitives Echolot bis auf 10 cm 
Entfernung. Bemerkenswert ist die zu dieser tiefen Frequenz passende Hörschärfe. 
Arius felis hört im Bereich 50 - 1000 Hz, mit bester Hörschärfe bei 100 - 200 Hz 
(Popper & Tavolga, 1981). Diese Werte liegen weit unter denen von Ameiurus 
nebulosus (Ictaluridae) mit 100 - 4000 Hz bzw. 600 - 800 Hz (Poggendorf, 1952, 
Weiss et al., 1969). In Feldstudien wurde gezeigt, daß A. felis besonders nachts Laute 
ausstößt, die dem Zusammenhalt der Gruppe dienen sollen. In Laborversuchen waren 
die Laute in Gesellschaft kürzer als unter Isolation. Ungestört gaben die Welse leise, 
20 - 40 ms kurze Grunzer von sich; bei Störung dagegen wurden laute, 100 - 150 ms 
lange Geräusche mit einer Grundfrequenz von 150 Hz registriert (Tavolga, 1960, 
1976). Bei Langzeituntersuchungen an A. felis fand Breder (1968), daß die Welse von 
April bis Oktober mit einer Sommerpause im Juli/August Laute erzeugten, haupt- 
sächlich von 17 - 23 Uhr bei einer Wassertemperatur von 23 - 31°C. Die Trommel- 
aktivität von A. seemani zeigte im Labor einen anderen Tag-Nacht-Rhythmus. Sie 
begann vor der Lichtperiode und war in den Morgenstunden am höchsten; nachts 
wurden kaum Laute registriert. Eine Funktion der Laute für den Zusammenhalt der 
Gruppe oder für die Orientierung im Dunkeln ist daher unwahrscheinlich. Bei 
Fischarten außerhalb der Welse wurden ebenfalls Dämmerungslaute (morgens und 
abends) beschrieben, deren biologische Bedeutung jedoch unbekannt ist (Winn, 1964; 
Mehickvet al 11999). 

Im Gegensatz zu den Untersuchungen an A. felis (Tavolga 1960) unterbrach A. 
seemani seine Trommelmuskelaktivität bei Störungen kurz, und die Lautdauer (ca. 
200 ms) blieb vor und nach Störungen nahezu unverändert. Die Grundfrequenz der 
Laute lag mit 140 - 200 Hz ım Bereich der von A. felis bekannten Kontraktions- 
frequenz des Trommelmuskels (Fusionsgrenze < 400 Hz). Die anatomischen Über- 
einstimmungen zwischen beiden Species deuten darauf hin, daß auch bei A. seemani 
die Grundfrequenz der Laute der Kontraktionsfrequenz des Protractors entspricht. Die 
Pausen zwischen den Lauten eines isolierten A. seemani schwankten in Ruhe und bei 
Störung. Arius seemani besitzt einen Springfederapparat wie A. felis (Tavolga 1962). 
Einen solchen Apparat führen auch die Auchenipteridae, Bagridae, Doradidae, 
Malapteruridae, Mochokidae und Pangasiidae (Müller, 1842; Bridge & Haddon, 
1893; Chranilov, 1929; Tavolga, 1962). Sörensen (1884) sowie Bridge & Haddon 
(1894) wiesen bereits darauf hin, daß der Springfederapparat der Doradidae der Laut- 
erzeugung dient, wie von Kastberger (1977, 1978) bestätigt. Entsprechendes gilt für 
die Mochokidae (Abu-Gideiri & Nasr, 1973). Der Springfederapparat und der direkte 
Trommelmechanismus (bei Pimelodidae) haben sich möglicherweise bei verschie- 
denen Welsfamilien unabhängig voneinander entwickelt (Alexander, 1965). Der 
Funktionsmechanismus des Springfederapparats läßt sich folgendermaßen beschrei- 
ben: Durch Kontraktion des Trommelmuskels (Protractors) werden der „Müller’sche 
Ramus“ (Tavolga, 1962) und der Transversalfortsatz des 4. Wirbels (Knochenplatte) 
craniad bzw. dorsad bewegt. Weil der Müller’sche Ramus über Bindegewebe mit der 
Tunica externa der vorderen Schwimmblasenkammer verwachsen ist, wird die 
Schwimmblase bei seiner Bewegung mitgezogen. Entspannt sich der Protractor, 


194 ANDREAS HEYD & WOLFGANG PFEIFFER 


schnellen Knochenplatte und Müller’scher Ramus wieder zurück. Die vordere 
Schwimmblasenkammer wird komprimiert und ihr Gasinhalt in die hintere Kammer 
gepreBt. Die Schwimmblase ist durch Transversalsepten unterteilt. Das an den 
Septenrändern und dem Diaphragma vorbeiströmende Gas versetzt diese in Schwin- 
gung. Die Schwimmblasenwand wird zum Vibrieren angeregt und ihre Schwingung 
durch die trommelnden Knochenelemente verstärkt (vergleichbar einem Schlagzeug). 
Die Schwimmblase dient als Resonanzkörper (Trommel). Schwingungsgeneratoren 
sind der Trommelmuskel und die knöchernen Elemente. Die Synchronisation der 
beiden Trommelmuskeln, rechts und links am jeweiligen Müller’schen Ramus, wird 
zurückgeführt auf die polyaxonale und multiple Versorgung ihrer Fasern durch den 
zweiten Spinalnerven (Schneider, 1967). 

Aspredinidae: Die Stridulationslaute von Bunocephalus spec. wurden von 
Winn (1964) analysiert und von Pfeiffer & Eisenberg (1965) an einigen Species 
gehört. Die Kontraktionsdauer der Muskelpotentiale von Bunocephalus spec. ent- 
spricht mit 100 ms der Lautlänge (Gainer, 1967). Die Lautdauer von Dysichthys 
coracoideus ist mit 80 ms nur wenig kürzer. Wie Bunocephalus spec. striduliert D. 
coracoideus bei Abduktion und Adduktion. 

Auchenipteridae: Sie besitzen neben der Brustflossenstridulation auch einen 
Springfederapparat (Müller 1842). Die Trommellaute von Trachycorystes spec. haben 
eine Grundfrequenz von 120 Hz und können einige Sekunden andauern (Kastberger, 
1978). Weitere Daten zu Schwimmblasen- und Stridulationslauten finden sich bei 
Kaatz (1999). 

Bagridae: Leiocassis micropogon erzeugt „knurrende“, L. siamensis und L. 
poecilopterus produzieren „krächzende“ Laute. L. brashnikowi gibt „einzelne Töne“ 
von sich, und beim Paarungsspiel von Mystus vittatus entstehen „zwitschernde“ Laute 
(Riehl & Baensch, 1983; Franke, 1985; Sterba, 1990), wobei der Mechanismus der 
Lauterzeugung nicht genannt wird. Mystus vittatus und M. gulio stridulieren bei 
Abduktion und Adduktion der Brustflossen. Mit den „Quietschlauten“ von Hemi- 
bagrus nemurus gibt es eine weitere, nämlich pneumatische Weise der Lautent- 
stehung. Agassız (1852) hörte bei einem Wels Geräusche, die anscheinend durch 
Luftausstossen aus der Schwimmblase hervorgerufen wurden. Bei Synodontis schall 
ist ein „Atemgeräusch“ vernehmbar, das vermutlich mit der Schwimmblase zusam- 
menhänst (Dufossé 1874). Sörensen (1895) beschrieb ein katzenähnliches Fauchen 
(„not unlike the hissing of a cat”) beim Zitterwels Malapterurus und führte dies auf 
die zwischen hinterer und vorderer Schwimmblasenkammer durch einen dünnen 
Gang streichende Luft zurück. Kaatz (1999) beobachtete Lauterzeugung durch 
Luftausstoßen bei 8 Welsfamilien. 

Callichthyidae: Callichthys callichthys läßt bei der Brutpflege „grunzende“ 
Laute hören (Riehl & Baensch, 1983). Hoplosternum thoracatum erzeugt bei 
Geschlechtsreife im Wasser Stridulationslaute (Mayr 1987). Für Dianema urostriata 
wurden Stridulationslaute nur bei Abduktion nachgewiesen, wie von H. thoracatum 
bekannt. Bei Corydoras paleatus stridulieren beide Geschlechter beim Fang, die 
Männchen auch bei Balz und Paarung (Pruzsinsky & Ladich, 1998). 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 195 


Clariidae / Heteropneustidae: Stridulationsgeräusche wurden für Hetero- 
pneustes fossilis beschrieben. Die Laute entstehen zum einen durch Bewegung der 
Brustflossen, zum anderen durch Reiben der Pharyngealzähne am Mundboden. Es 
wird vermutet, daß die Laute der Warnung und Abwehr von Angreifern sowie 
während der Fortpflanzungsperiode der Schwarmbildung dienen (Agrawal & Sharma 
1965). Diese Species, früher zu den Clariidae gezählt, wird heute den Hete- 
ropneustidae zugeordnet (Burgess, 1989; Teugels, 1996). Über die Lauterzeugung der 
Clariidae gibt es kaum Zeugnisse: Tennent (1859; zitiert in Sörensen, 1895, p. 115f.) 
berichtet über Aussagen von Fischern, nach denen ein „Magoora“ genannter Fisch in 
einem See nahe Colombo bei Störung im Wasser „Grunzlaute‘“ hervorbringt. Nach 
Day (1881) handelt es sich dabei um einen Wels der Gattung Clarias, wahrscheinlich 
um Clarias magur (,,Mah-gur* oder „Magurah“, Day, 1958), einem Synonym von 
Clarias batrachus (Eschmeyer, 1998). Der Laut soll durch Schwimmblasenluft ent- 
stehen, die über den Ductus pneumaticus entweicht (Sörensen, 1895). Es könnte sich 
aber auch um Stridulationslaute handeln, da viele Clariidae kräftige Brustflossen- 
stacheln besitzen, deren Struktur derjenigen stridulierender Species ähnelt (Nawar 
1954). 

Doradidae: Acanthodoras spinosissimus „grunzt“ (Innes 1956), „Knurrt‘ (Ster- 
ba 1956), „krächzt‘ und gibt einen lang anhaltenden Quarrlaut von sich, der aus einer 
Folge von Einzellauten besteht (Villwock, 1960). Amblydoras hancocki erzeugt 
knurrende Töne, und Platydoras costatus stößt beim Fangen „quakende“ Laute aus 
(Sterba 1990). Die Lauterzeugung in Abhängigkeit von der Bewegung der Brust- 
flossen wurde erstmals von Pfeiffer & Eisenberg (1965) experimentell untersucht, 
durch gleichzeitige Film- und Tonbandaufnahmen von Agamyxis albomaculatus, 
Amblydoras hancocki und Platydoras costatus. Die Laute entstehen bei Abduktion 
und Adduktion, wobei die Abduktionslaute länger sind als die Adduktionslaute. Die 
an Agamyxis flavopictus erhobenen Befunde bezüglich Laut- und Pausenlängen 
entsprechen den Ergebnissen von Pfeiffer & Eisenberg (1965). Dagegen fand Ladich 
(1997), daß sich Ab- und Adduktionslaute bei Agamyxis pectinifrons bzw. Platydoras 
costatus ın der Länge nicht unterscheiden. Die Dauer der Stridulationslaute liegt bei 
den beiden Species A. flavopictus und A. pectinifrons mit ca. 100 ms im gleichen 
Bereich. A. flavopictus wird 1m Handel oft als A. pectinifrons bezeichnet (Franke 
1985). Hassar orestis kann ebenfalls stridulieren und ist durch seine abgespreizten 
Flossenstacheln sogar vor Piranhas geschützt (Markl, 1968). Neben der Brustflossen- 
stridulation besitzen die Doradidae einen indirekten Trommelmechanismus, den 
“Springfederapparat” (Müller, 1842; Sörensen, 1884, 1895; Bridge & Haddon, 1894; 
Dorn, 1976; Kastberger, 1977, 1978). Die biologische Bedeutung der durch die 
Brustflossenstridulation erzeugten Laute sehen Pfeiffer & Eisenberg (1965) im 
„Schutz oder der Verteidigung“, in der Warnung eines potentiellen Beutegreifers vor 
einem nur schwer zu verschlingenden, gefährlichen Happen. Die Gefährlichkeit ist 
gegeben durch die Sperrmechanismen der Stacheln von Brustflossen und Rücken- 
flosse (Brousseau, 1976), in Verbindung mit ihrer großen mechanischen Festigkeit 
(Schaefer, 1984) und möglichen Giftigkeit (Birkhead, 1972), die sogar für den 
Menschen tödlich sein kann (Nelson, 1994). 


196 ANDREAS HEYD & WOLFGANG PFEIFFER 


Ictaluridae: Den Bau des Schultergürtels von /ctalurus nebulosus beschreibt 
Brousseau (1976). I. nebulosus striduliert bei Abduktion und Adduktion, wodurch die 
Anzahl der aggressiven Auseinandersetzungen zwischen dem Revierinhaber und 
einem Eindringling herabgesetzt wird (Rigley & Muir, 1979). Dagegen striduliert 
Ictalurus punctatus nur bei Abduktion, meistens mit der rechten Brustflosse (Fine et 
al., 1996, 1997). Daß /ctalurus nebulosus nach Lundberg (1992) zur Gattung 
Ameiurus gehört, könnte den Unterschied im Stridulationsmechanismus erklären. 
Jedoch sind nach Eschmeyer (1998) beide Gattungsnamen für diese Species gültig. 
Die Sperrmechanismen des Rückenflossenstachels und der Brustflossenstacheln 
schützen /. melas weitgehend davor, von Reihern und Rothalstauchern gefressen zu 
werden, die ihre Beute nicht zerteilen, nicht aber vor Seeadlern, die sie zerlegen. Die 
beiden kleineren Vogelarten fressen überwiegend Barsche (Forbes, 1989). 

Loricariidae: Plecostomus spec. striduliert und besitzt ein ähnliches Brust- 
flossenskelett wie die Doradidae und Mochokidae (Pfeiffer & Eisenberg, 1965). 
Peckoltia pulcher und Glyptoperichthys gibbiceps stridulieren nur bei Abduktion. Bei 
Pterygoplichthys spec. beträgt die Lautdauer 250 ms, die Grundfrequenz 500 Hz, die 
Pausendauer 400 - 600 ms. Große Exemplare zeigen eine kürzere Laut- und Pausen- 
dauer und ein höher liegendes Frequenzspektrum als kleine (Schachner, 1977). 
Flossenstacheln und Schultergürtel von Prerygoplichthys spec. sind von enormer 
mechanischer Stabilität (Schaefer, 1984). 

Mochokidae: Synodontis spec. ist bereits vor 170 Jahren als erster durch 
Stridulation Laute erzeugender Wels in die Literatur eingegangen (Geoffroy Saint- 
Hilaire, 1829). Aus Daget et al. (1986) geht hervor, daß es sich hierbei um Synodontis 
schall (Bloch & Schneider, 1801) handelt. Diese Species striduliert bei Abduktion 
und Adduktion der Brustflosse (Müller, 1857, Dufossé, 1874) wie S. nigrita und S. 
nigriventris (Pfeiffer & Eisenberg, 1965), S. ocellifer und S. schoutedeni. Die Be- 
funde an S. ocellifer und S. schoutedeni entsprechen in jeder Beziehung (Lautmecha- 
nismus, Laut- und Pausendauer, Struktur der Sonagramme und Frequenzschwer- 
punkte) den vor 35 Jahre vom Zweit-Autor erhobenen (Pfeiffer & Eisenberg, 1965). 
Alle gepriiften Mochokidae stridulieren schneller als die Doradidae, weshalb die 
Dauer ihrer Laute und Intervalle kürzer ist. Ladich (1997) konnte keinen signifikanten 
Unterschied in den Lautlängen zwischen diesen beiden Familien finden, was für die 
Abduktionslaute knapp zutrifft (p = 0.08). Die Adduktionslaute der Doradidae waren 
länger als die der Mochokidae. Zur Klärung dieser Frage sollten weitere Species 
beider Familien untersucht werden. Die Mochokidae besitzen außer der Brustflossen- 
Stridulation auch einen Springfederapparat (Sörensen, 1884, 1895; Bridge & Haddon, 
1893). S. schall striduliert und trommelt. Er soll Laute erzeugen bei Verfolgung, 
„Schmerz“, aggressivem Verhalten, Laichverhalten und als Reaktion auf Berührungs- 
reize (Abu-Gideiri & Nasr, 1973). 

Pangasiidae: Pangasius sutchi erzeugt Stridulationsgeräusche (Kaatz, 1999). 

Pimelodidae: Bau und Funktion des Stridulationsapparates und des Trommel- 
mechanismus wurden von Schachner (1977) an Pimelodus spec. untersucht. Schach- 
ner & Schaller (1982) zeigen, daß es sich dabei um Rhamdia sebae handelt und 
beschreiben Defensivgeräusche (Stridulation), Drohlaute (Intervalle 60 - 70 ms) und 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 197 


„Störungslaute“ (Trommellaute wie die Drohlaute, jedoch mit kürzeren Intervallen 
von nur 10 - 20 ms und höherer Frequenz). Die Stridulationsgeräusche dauern 40 - 85 
ms, im Mittel 54 ms; ihr Energieschwerpunkt liegt bei 2 KHz (Schachner 1977). Die 
Stridulationsgeräusche von Brachyrhamdia meesi entstehen nur bei Abduktion der 
Brustflosse, wie diejenigen von R. sebae. Auch bei dem von Kratochvil er al. (1980) 
und Kratochvil & Völlenkle (1981) untersuchten Pimelodus spec. dienen die 
Stridulationslaute bei Rivalenkämpfen als Warnlaute, zur Revierverteidigung und bei 
interspecifischen Attacken. Stoß und Biß werden mit Stridulationsgeräuschen beant- 
wortet. Akustische Drohsignale können Fluchtreaktionen auslösen. Die Trommellaute 
dienen der innerartlichen Verteidigung, die Stridulationslaute der Warnung. Den 
Feinbau der Trommelmuskeln beschreibt Dorn (1976). Die für die Lauterzeugung 
verantwortlichen Motoneuronen identifizierten Ladich & Fine (1994) an Pimelodus 
blochi und P. pictus. Drei Äste des N. occipitalis und die beiden ersten Spinalnerven 
versorgen das Trommelorgan, ein rostraler Ast des N. occipitalis und die beiden 
ersten Spinalnerven die Stridulationsmuskeln (Abduktor und Adduktor). Pimelodus 
pictus zeigte Drohverhalten gegenüber Artgenossen durch Spreizen seiner Flossen 
und Kiemendeckel, gleichzeitig Trommellaute erzeugend, wie von Schachner & 
Schaller (1982) fiir Rhamdia sebae geschildert. Auch die Strukturen des Trommel- 
mechanismus von P. pictus entsprechen weitgehend der Beschreibung für R. sebae. In 
beiden Fällen handelt es sich um einen direkten Trommelmuskelapparat. Der 
anteriore Teil des transversalen Fortsatzes vom 4. Wirbel bildet einen Wulst, an dem 
der Trommelmuskel ventral entspringt. Bei beiden Species setzt der Trommelmuskel 
S-förmig an der ventralen Schwimmblasenwand an, wobei er bei P. pictus etwas 
weiter tiber den caudalen Rand der vorderen Schwimmblasenkammer hinwegzieht als 
bei R. sebae. Die Kammerung des posterioren Schwimmblasenteils ist verschieden: 
bei R. sebae treten zahlreiche Transversalsepten auf, bei P. pictus nur drei. Laut- 
bildung und Funktion der Transversalsepten einschließlich des Diaphragmas der 
Schwimmblase wurden bereits von Sörensen (1884, 1895) beschrieben. Durch 
Kontraktion des Trommelmuskels wird die vordere Schwimmblasenkammer kompri- 
miert und ihr Gasinhalt in die hintere Kammer gepreßt. Beim Entspannen des 
Muskels wird das Gas wieder zurückgesogen. An den Rändern der unvollständigen 
Transversalsepten vorbeistreichend versetzt es diese in Vibration und regt die gesamte 
Schwimmblase zum Schwingen an. Der kontraktierende Trommelmuskel erzeugt das 
Geräusch, das durch die Schwingungen von Septen und Schwimmblase verstärkt 
wird. Sörensen (1895) schnitt winzige Löcher in die Schwimmblasenwand, wodurch 
der Laut an Intensität verlor. Doch war der Laut sogar nach Entfernen der Schwimm- 
blase noch schwach zu hören. Dagegen war nach Durchtrennung des paarigen Trom- 
melmuskels kein Geräusch mehr wahrzunehmen. Bei R. sebae sind im posterioren 
Schwimmblasenabschnitt nur die Septen beweglich, die Schwimmblasenwand ist starr 
(Schachner & Schaller, 1982). Außer den Septen wird vor allem der vordere 
Schwimmblasenabschnitt zum Schwingen angeregt, so daß die Vibrationen beidseitig 
über die lateralen Fenster in der Muskulatur direkt an das umgebende Wasser 
abgegeben werden können. Bei der Präparation der Schwimmblase von P. pictus war 
auffallend, daß ihr posteriorer Teil stabiler ist als ihr anteriorer. Wurde die 


198 ANDREAS HEYD & WOLFGANG PFEIFFER 


Schwimmblase lateral eröffnet, fiel die vordere Kammer zusammen, während die 
hintere stabil blieb. Der posteriore Schwimmblasenteil wird durch die Transver- 
salsepten und die festere Wand gestützt. Tavolga (1962) reizte Trommelmuskeln von 
Bagre marinus elektrisch: die zur Dauerkontraktion notwendige Frequenz beträgt 500 
Hz. Die Fasern der Trommelmuskeln von P. pictus gehören zu den schnell zuckenden 
und sind dicht kapillarisiert. Ontogenetisch leiten sich die Trommelmuskeln von 
Myotom-Muskulatur ab, die in Richtung Peritoneum gewandert ist und sich diesem 
aufgelagert hat. Das Peritoneum ist an der Ansatzstelle des Trommelmuskels an der 
Schwimmblase mit deren Tunica externa verwachsen (Alexander, 1965). Der direkte 
Trommelmuskelapparat deutet darauf hin, daß P. pictus Trommellaute erzeugt. 
Pimelodella gracilis besitzt wie Pimelodus pictus beidseits je ein Fenster in der 
Muskulatur, Transversalfortsätze des 4. und 5. Wirbels und eine gut ausgebildete 
Schwimmblase. Ein Trommelmuskel konnte bei P. gracilis nicht gefunden werden. 
Da nur juvenile Weibchen untersucht wurden, ist nicht auszuschließen, daß der 
Trommelmuskel bei diesen Fischen entweder noch nicht entwickelt war oder nur bei 
Männchen existiert. Die folgenden Tatsachen sprechen gegen einen Trommelapparat 
bei P. gracilis: ein derart mächtiger Trommelmuskel, wie ihn P. pictus und Arius 
seemani haben, wird vermutlich bereits in einem frühen Entwicklungsstadium an- 
gelegt; die untersuchten juvenilen P. pictus besaßen bereits einen Trommelmecha- 
nismus. Ferner spricht die Form der Schwimmblase von P. gracilis gegen eine 
Funktion als Lautorgan. Schließlich fehlen der hinteren Schwimmblasenkammer 
Transversalsepten, die nach Sörensen (1895) für die Lauterzeugung notwendig sind. 
Die posteriore Kammer ist durch ein Longitudinalseptum zweigeteilt. Das Gas in der 
Schwimmblase hat nur die Möglichkeit, an den Diaphragma-Rändern entlang in eine 
dieser beiden hinteren Kammern zu gelangen. Zudem ist die vordere Kammer von 
P. gracilis die kleinere, im Gegensatz zu P. pictus. Es könnte also nur ein geringes 
Gasvolumen bewegt werden, um das Diaphragma und die Schwimmblasenwände 
zum Schwingen anzuregen. Die beiden lateralen Fenster in der Muskulatur und die 
knöchernen Elemente des 4. und 5. Wirbels sind für das Hören wichtig (Alexander, 
1965). Diese Fenster verringern den Widerstand der Körperwand, so daß die 
Schwimmblase Schallschwingungen fast ungedämpft aufnehmen kann, um sie über 
die ihr craniad-dorsad aufliegenden Weberschen Knöchelchen an das Innenohr 
weiterzuleiten. 

Plotosidae: Burgess (1989) zitiert Beschreibungen von Plotosus lineatus, in 
denen diese Species auch ,,bumblebee catfish“ (Hummelwels) genannt wird. Dem- 
nach soll ein Summen hörbar sein, sobald man diese Welse aus dem Wasser fischt. 
Auch bei Störungen im Wasser wurden solche Geräusche vernommen. Da ein 
Schwimmblasenmechanismus bei dieser Species nicht beschrieben wurde, könnte es 
sich um Stridulationslaute handeln. 

Siluridae: Stridulationsgeräusche wurden für diese Familie bisher nicht 
beschrieben. Bei Hawkins (1986) findet sich aber folgende Bemerkung: „Some 
catfish of the family Siluridae produce a squeak when the enlarged pectoral spines are 
moved.“ Da eine Quellenangabe fehlt, handelt es sich möglicherweise um eine 
Verwechslung mit anderen Welsfamilien. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 199 


Sisoridae: Im Gegensatz zu allen anderen Welsfamilien stridulieren die beiden 
bisher untersuchten Sisoridae nicht mit den Brustflossen, sondern mit der Rücken- 
flosse, wie von Haddon (1881) an Gagata gagata entdeckt und von Mahajan (1963) 
an Sisor rhabdophorus bestätigt. Die Lauterzeugung geschieht bei Abduktion und 
Adduktion der Rückenflosse und dient vermutlich dazu, Beutegreifer vor der Gefähr- 
lichkeit oder Ungenießbarkeit des Fisches wegen seiner kräftigen Brustflossen- 
stacheln und seines Rückenflossenstachels zu warnen. 

Trommelmechanismen sind nach der Stridulation die häufigste Form der 
Lauterzeugung. Die Frage, ob die Schwimmblase für die Lauterzeugung notwendig 
ist, wird unterschiedlich beantwortet, obwohl es auch lauterzeugende Fische ohne 
Schwimmblase gibt. Während die Schwimmblase schon früh als hydrostatisches 
Organ erkannt wurde (Boyle, 1670, 1675; Ray, 1675), ist ihre große Bedeutung für 
Hörvermögen, Weberschen Apparat, Lauterzeugung und akustische Kommunikation 
erst seit dem 19. und 20. Jahrhundert bekannt: (1) Hörvermögen (anatomisch): 
Weber, 1820; Reissner, 1859; Bridge & Haddon, 1893; Chranilov, 1929 und Chardon, 
1968; (physiologisch) von Frisch & Stetter, 1932; Poggendorf, 1952 und Weiss et a/., 
1969; (Reviews) Harden Jones & Marshall, 1953; Alexander, 1964, 1965, 1966; 
Popper & Fay, 1973, 1993; Rogers et al., 1988 und Schellart & Wubbels, 1998; (2) 
Lauterzeugung (anatomisch): Müller, 1857; Dufossé, 1874; Moreau, 1876; Sörensen, 
1884, 1895 und Bridge & Haddon, 1889; (Reviews) Harden Jones & Marshall, 1953; 
Schneider, 1961, 1967 und Demski, et al., 1973; (3) akustische Kommunikation 
(Reviews): Tavolga, 1960, 1971a, 1977, Fine, et al., 1977; Myrberg, 1981; Hawkins 
& Myrberg, 1983; Michelsen, 1983; Urick, 1983 und Tyack, 1998. 

Die Schwimmblase ist bei der Lauterzeugung durch Trommelmechanismen 
der Siluroidei beteiligt (Miiller, 1857; Tavolga, 1962; Ladich, 1997). Durch ihre 
Gasfillung fungiert sie als Impedanzwandler zwischen den Trommelmuskeln und 
dem Wasser und verstärkt so die erzeugten Schwingungen. Es wird diskutiert, ob 
dabei diejenigen Frequenzen am besten verstärkt werden, die ihrer Resonanzfrequenz 
entsprechen (Demski ef al., 1973). Kaatz (1995) fand bei mehreren Species der 
Auchenipteridae und Doradidae Korrelationen zwischen der Schwimmblasengröße 
und der Hauptfrequenz der erzeugten Laute. Bei Fischarten außerhalb der Siluroidei 
wurde festgestellt, daß größere Exemplare tiefere Laute hervorbringen. Dies wird auf 
veränderte Resonanzeigenschaften der Schwimmblase zurückgeführt (Demski et al., 
1973), da die Resonanzfrequenz mit zunehmendem Volumen der Schwimmblase 
sinkt (Urick, 1983). Es muß dabei berücksichtigt werden, daß die Schwimmblase von 
verschiedenen Geweben umgeben ist, die Schwingungsdämpfungen bewirken und 
ihre Resonanzeigenschaften beeinflussen. Bei Stridulationsgeräuschen ist die Betei- 
ligung der Schwimmblase umstritten: in der Lautstärke besteht außerhalb des Wassers 
bei intakten Doradidae und Mochokidae einerseits und Exemplaren mit eröffneter 
Schwimmblase andererseits kein Unterschied (Pfeiffer & Eisenberg, 1965). Zur 
selben Ansicht kamen Fine er al. (1997) bei Untersuchungen an /ctalurus punctatus: 
hier spielt die Schwimmblase bezüglich Frequenzspektrum und Amplitude der Stridu- 
lationslaute zumindest keine Hauptrolle, während eher der Schultergürtel auf die 


200 ANDREAS HEYD & WOLFGANG PFEIFFER 


Frequenzverteilung Einfluß nimmt. Andererseits ergaben Untersuchungen an Cory- 
doras paleatus, daß die Hauptfrequenz der Stridulationslaute mit zunehmender 
Körpermasse abnimmt (Pruzsinsky & Ladich 1998). Es wäre möglich, daß hier 
veränderte Resonanzeigenschaften der Schwimmblase eine Rolle spielen. Ob stridu- 
lierende Fische die Resonanzfrequenz ihrer Schwimmblase modulieren, wurde bisher 
nicht untersucht (Zelick er al., 1999). 


LAUTERZEUGUNG UND PHYLOGENIE 


Wichtige Fragen zu den Welsen (Berg, 1958; Nelson, 1976; Burgess, 1989) 
sind ungelöst. Über die Phylogenie und das Natürliche System gibt es verschiedene, 
zum Teil konträre Meinungen (Eigenmann & Eigenmann, 1890; Regan, 1911, Green- 
wood et al., 1966; Rosen & Greenwood, 1970; Roberts, 1973; Lauder & Liem, 1983; 
Fink & Fink, 1981, 1996; Schaefer, 1987). Die Klassifizierung in Familien sowie die 
Anzahl der Gattungen und Species hat sich laufend verändert (Tab. 3; Mo, 1991; 
Nelson, 1994; Teugels, 1996). Auch die Verbreitungsgeschichte der Welse ist unklar: 
diskutiert werden primäre Zentren in Südamerika und Südostasien, ein sekundäres 
Zentrum in Afrika (Gosline, 1975) oder Entstehung in Südamerika bzw. Südostasien 
(Novacek & Marshall, 1976; Briggs, 1979). Die Fossilgeschichte ist ebenfalls 
unbefriedigend; vermutlich war die interkontinentale Verbreitung schon vor dem 
Eozän abgeschlossen (Gosline, 1975). Innerhalb der Siluroidei gibt es so viel Parallel- 
Evolution, daß die phylogenetischen Zusammenhänge bei unserem gegenwärtigen 
Wissensstand unklar bleiben (Alexander, 1965; Teugels, 1996; de Pinna, 1998). 

In ihrer Kommunikation zeigen die Welse ein vielfältiges Bild. Sie verfügen 
über mindestens 6 verschiedene Möglichkeiten der Lauterzeugung (Tab. 3): (1) 
Stridulation durch Abduktion der Brustflossen (Ariidae, Callichthyidae, Ictaluridae, 
Loricariidae, Pimelodidae), (2) Stridulation durch Abduktion und Adduktion der 
Brustflossen (Aspredinidae, Bagridae, Doradidae, Ictaluridae, Mochokidae), (3) Stri- 
dulation mit der Rückenflosse (Sisoridae), (4) pneumatische Mechanismen durch 
Luftausstoßen über die Kiemenspalten (Hemibagrus nemurus), (5) direkter Trommel- 
mechanismus (Pimelodus pictus) und (6) indirekter Trommelmechanismus oder 
Springfederapparat (Arius seemani). Innerhalb der Familien und Gattungen können 
verschiedene Mechanismen auftreten oder fehlen. Bei den Loricariidae gibt es neben 
stridulierenden Species nichtstridulierende, sogar innerhalb derselben Unterfamilie 
(Ancistrinae; Schaefer 1987). Doch sind die Verwandtschaftsverhältnisse innerhalb 
der Loricariidae ungeklärt. Nach Untersuchungen mit Hilfe der rRNA-Sequenzierung 
sind alle Unterfamilien außer den Loricasiinae paraphyletisch (Montoya-Burgos et al. 
1998). 

Bei den Bagridae stridulieren Mystus gulio und M. vittatus, während A. 
nemurus mit Hilfe der Kiemenspalten quietscht. Auch der Stridulationsmechanismus 
kann in derselben Familie wechseln (Ictaluridae: Ameiurus, Ictalurus). Innerhalb 
derselben Species treten stridulierende und nichtstridulierende Individuen auf, wobei 
der Anteil der stridulierenden Exemplare unterschiedlich ist. Die einzelnen Parameter 
der Geräusche wie Häufigkeit, Wiederholungsrate, Laut- und Intervalldauer, Laut- 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 201 


stärke oder Frequenzanteile variieren bei Familien, Arten, Angehörigen derselben 
Species und sogar bei ein und demselben Individuum. Die an einer Species erhobenen 
Befunde lassen sich weder für eine Familie noch für eine Gattung verallgemeinern. 

Die weite Verbreitung der Lauterzeugung durch Stridulation der Brustflossen 
sowie die weitgehend ähnliche strukturelle Ausbildung des Stridulationsapparates 
deuten daraufhin, daß dieser Mechanismus bei manchen Familien sekundär teilweise 
oder ganz verlorengegangen ist. Die Stridulationsmechanismen Abduktion und 
Adduktion der Brustflossen sind sowohl gemeinsam als auch allein (nur Abduktion) 
in allen größeren Verwandtschaftsgruppen zu finden (de Pinna, 1998; Tab. 3). Ent- 
sprechende Untersuchungen der Basisgruppen der Welse (Diplomystidae, Cetopsidae 
u.a.) liegen bisher nicht vor. Stridulationslaute treten außer bei Störungen auch beim 
Fortpflanzungsverhalten mancher Species auf (Mayr, 1987; Pruzsinsky & Ladich, 
1998; Kaatz & Lobel, 1999). Es gibt Hinweise, daß der Lauterzeugungsapparat bei 
diesen Species sekundär abgewandelt wurde (Kaatz & Stewart, 1997). 

Die Schwimmblasenmechanismen sind unterschiedlich gestaltet, doch nicht 
familiencharakteristisch, wie Ladich (1997) meint. Bei den Pimelodidae besitzt 
Pimelodus pictus einen Trommelmechanismus, Pimelodella gracilis nicht. Es könnte 
sich dabei um einen Unterschied zwischen Unterfamilien handeln, da Pimelodus zu 
den Pimelodinae, Pimelodella aber zu den Rhamdiinae (Teugels, 1996) gestellt wird. 
Die letztgenannte Gruppe wird von de Pinna (1998) als Heptapterinae bezeichnet, 
deren Phylogenie ungeklärt ist. Doch treten auch innerhalb derselben Unterfamilie 
(Pimelodinae, Calophysus-Gruppe) Species mit und ohne Schwimmblasenmuskeln 
auf (Stewart, 1986; de Pinna, 1998). Weder bei den Basisgruppen Diplomystidae, 
Cetopsidae noch bei der nächsthöheren, 12 Familien umfassenden, Gruppe (Lorica- 
ruidae-Callichthyidae-Aspredinidae-Trichomycteridae u.a.; de Pinna, 1998) ist bisher 
ein Trommelmuskelapparat gefunden worden. 


LAUTERZEUGUNG UND SCHRECKREAKTION 


Die Schreckreaktion auf Schreckstoff aus der Haut von Artgenossen wurde 
von Karl von Frisch (1938) an der Elritze Phoxinus phoxinus (L.) (Cyprinidae) ent- 
deckt und später auch an fünf anderen Species der Cyprinidae nachgewiesen. Haut- 
extrakt von 20 weiteren Species einheimischer Cyprinidae und zwei Species der 
Cobitidae löste bei Elritzenschwärmen ebenfalls eine Schreckreaktion aus. Dagegen 
waren die Haute von allen Fischen, die nicht zu den Cyprinidae und Cobitidae 
gehören, darunter der nordamerikanische Zwergwels Ameiurus nebulosus, unwirk- 
sam. Der einzige Versuch mit diesem Wels endete negativ (von Frisch, 1941a,b). 
Zwei Schüler von Karl von Frisch setzten die von ihm begonnenen Versuche über die 
Verbreitung der Schreckreaktion fort: F. Schutz (1956) und W. Pfeiffer (1960 bis 
heute). Nach Schutz (1956) ist die Schreckreaktion unter den Cyprinidae allgemein 
verbreitet, selbst wenn es sich nicht um ausgesprochene Schwarmbildner handelt. 
„Die gleiche Reaktion findet sich auch bei kleineren, friedlich lebenden und 
schwarmbildenden Characidae.“ Auch alle vier Versuchsgruppen von Ameiurus nebu- 
losus zeigten in acht Versuchen immer ein positives Ergebnis, das Schutz (1956) 


202 ANDREAS HEYD & WOLFGANG PFEIFFER 


jedoch als unspezifische Reaktion „von mehr ausweichendem als schreckhaftem 
Charakter“ betrachtete: „Trotz der Höhe der Konzentration der arteigenen Haut- 
extrakte waren alle Reaktionen immer nur kurz und ohne nachhaltige Wirkung.“ 
Schutz (1956) war der Auffassung, daß eine Schreckreaktion bei den Zwergwelsen 
bemerkenswert wäre, weil die jüngeren Fische in Schulen zusammenhalten. Auch die 
kleinen Panzerwelse Corydoras paleatus (Callichthyidae) schwammen immer in einer 
Gruppe und zeigten auf ihren Hautextrakt „leicht schreckhaftes Verhalten.“ Ihre 
Reaktionsweise erzeugte bei Schutz (1956) jedoch den Eindruck, „daß es sich wie 
beim Zwergwels um eine unspezifische Reaktion handelt“. 

Pfeiffer (1960) zeigte in fünf Versuchen an zwei Schwärmen des indischen 
Glaswelses Kryptopterus bicirrhis (Siluridae) erstmals, daß auch Siluroidei eine 
Schreckreaktion besitzen, die derjenigen der Cypriniformes und Characiformes 
gleichzusetzen ist. Dieses Ergebnis hat sich später an dieser und anderen Species der 
Welse bestätigt (Pfeiffer, 1963a; Pfeiffer er al., 1986; Heyd & Pfeiffer, in Vor- 
bereitung) und beweist, daß die Schreckreaktion bei den Ostariophysen (sensu 
Sagemehl 1885) allgemein verbreitet ist. Die Entdeckung der Schreckreaktion bei den 
Gonorynchitormes (Pfeiffer 1967) fügt sich in das System von Rosen & Greenwood 
(1970) ein, wonach die Ostariophysen sensu Sagemehl (1885) als Otophysi mit den 
Anotophysi (alias Gonorynchiformes) als Ostariophysi zusammengefaßt werden. Alle 
analogen Alarm-Pheromon-Systeme von nichtostariophysen Fischen (Cottidae, Gobi- 
idae, Percidae) unterscheiden sich tiefgreifend von der Schreckreaktion der Ostario- 
physi (Smith, 1992). Nur die Ostariophysi (sensu Rosen & Greenwood, 1970) be- 
sitzen in ihrer Epidermis Schreckstoffzellen (Pfeiffer, 1960; Pfeiffer er al., 1971), 
denen der Schreckstoff entstammt (Pfeiffer, 1960, 1963a, 1967; Smith, 1973, 
1976a,b). 

Ob die Schreckstoffzellen noch weitere Funktionen haben, wie von Pfeiffer 
(1967, 1970) für solitäre, nachtaktive Welse vermutet, ist nicht entschieden. Um die 
besondere Situation der Siluroidei, was ihre Schreckreaktion anbetrifft, zu verstehen, 
werfen wir zunächst einen Blick auf die anderen, wesentlich intensiver untersuchten 
Ostariophysi: 

(1) Innerhalb der Cypriniformes wurde die Schreckreaktion bei allen mehr als 
53 bisher gepriiften Species aus fiinf Familien gefunden, mit alleiniger Ausnahme des 
blinden Höhlenfisches Caecobarbus geertsi. Sie existiert bei allen tagaktiven Cyprin- 
idae, auch bei bodenlebenden Species, sowie bei den Catostomidae, Cobitidae, 
Gyrinocheilidae und Homalopteridae. Auch die beiden räuberisch lebenden Cyprin- 
idae Leuciscus cephalus und Prychocheilus oregonense haben zumindest als Jung- 
fische eine Schreckreaktion (Pfeiffer 1977). 

(2) Innerhalb der Characiformes dagegen wurde die Schreckreaktion nur bei 
2/3 der mehr als 50 geprüften Species gefunden. Es gibt hier Familien bzw. Unter- 
familien, für welche die Schreckreaktion bisher nicht nachgewiesen werden konnte, 
nämlich die Chilodontidae, Lebiasinidae, Mylinae und Serrasalminae. Während das 
Fehlen der Schreckreaktion bei den räuberischen Piranhas verständlich ist, gilt dies 
nicht für die ihnen nahverwandten pflanzenfressenden und friedlichen Mylinae, es sei 
denn, man hält die Mylinae für Abkömmlinge der Serrasalminae. Beide, Serrasalm- 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 203 


inae und Mylinae, besitzen Schreckstoffzellen und Schreckstoff, doch fehlt ihnen im 
ZNS die Fähigkeit, auf den Schreckstoff zu reagieren. Entsprechendes gilt für den 
blinden Höhlensalmler Anoptichthys jordani (Pfeiffer, 1967, 1977). 

(3) Allen untersuchten Gymnotoidei fehlen Schreckstoffzellen und Schreck- 
reaktion (Pfeiffer, 1963a, 1967). 

(4) Bei allen 7 histologisch geprüften Gattungen der Anotophysi aus 4 Fami- 
lien wurden Schreckstoffzellen gefunden. Für die beiden im Verhaltensexperiment 
getesteten Gattungen wurde die Schreckreaktion nachgewiesen. Von besonderer 
Bedeutung ist die wechselseitige Wirksamkeit des Schreckstoffs zwischen Otophysi 
und Anotophysi (Pfeiffer, 1967). 

Von den ca. 33 Familien der Siluroidei wurden bisher 22 bezüglich ihrer 
Schreckstoffzellen untersucht. Bei 20 Welsfamilien wurden sie gefunden (Tab. 3), nur 
die Aspredinidae und Loricariidae führen keine (Pfeiffer, 1963c, 1977). Den Aspre- 
dinidae fehlen darüber hinaus sogar die Mauthner-Axone und damit der „Mauthner- 
Reflex“, mit dem jede Schreckreaktion beginnt (Göhner & Pfeiffer, 1996, Pfeiffer er 
al., 1986). Im Gegensatz zu den intensiv untersuchten Cypriniformes und Characi- 
formes wurden bisher nur verhältnismäßig wenige Species und Familien der Silur- 
oidei bezüglich ihrer Schreckreaktion geprüft (Pfeiffer, 1977), aus verständlichen 
Gründen, wie die Ergebnisse von Karl von Frisch (1938, 1941a, b) und Schutz (1956) 
zeigen. Die Schreckreaktion wurde bei Angehörigen der Clariidae, Ictaluridae, 
Mochokidae und Pimelodidae gesehen. Mit Sicherheit nachgewiesen, videotechnisch 
aufgezeichnet und analysiert wurde sie für Kryptopterus bicirrhis (Siluridae), Pan- 
gasius sutchi (Pangasiidae), Eutropiellus vanderweyeri (Schilbeidae), Corydoras 
paleatus (Callichthyidae) und Plotosus lineatus (Plotosidae) (Pfeiffer, 1960, 1963a, 
1977; Pfeiffer er al., 1986, Heyd & Pfeiffer, in Vorbereitung). 

Die Mehrzahl der Species und Familien der Welse sind abgeflachte Boden- 
fische mit hohem spezifischen Gewicht und unterständigem Maul, nachtaktiv und 
kleinäugig, mit Barteln als Trägern von Tast- und Geschmacksorganen und einem 
hervorragenden Hörvermögen. Sie sind meist solitär und wehrhaft durch kräftige 
Stacheln der Rückenflosse und der Brustflossen, die mit Sperrmechanismen aus- 
gestattet und als Stridulationsapparate prädestiniert sind. Die meisten Welse ernähren 
sich von kleinen Invertebraten, einige sind Räuber, andere Algen- und Detritusfresser. 
Ihre nächtliche Aktivität und ihr Bodenleben teilen sie mit den Gymnotoidei, mit 
denen die Aspredinidae und Loricariidae auch das Fehlen von Schreckstoffzellen 
sowie mehrere andere Welsfamilien den Besitz von Elektrorezeptoren („small pit 
organs”) gemeinsam haben. Ihre Lebensweise als solitäre nachtaktive Bodenfische 
unterscheidet die meisten Siluroidei und alle Gymnotoidei deutlich von den gewöhn- 
lich gesellig lebenden, tagaktiven, größtenteils pelagischen Cypriniformes, Characi- 
formes und Anotophysi. Die Welsarten mit der am deutlichsten ausgeprägten 
Schreckreaktion sind ebenfalls tagaktive Schwarmfische oder zumindest als Jung- 
fische in Gruppen lebend. Sie erinnern durch ihr Aussehen und ihre Lebensweise an 
dıe große Mehrzahl der Cypriniformes und Characiformes. Corydoras ist zwar boden- 
lebend, doch tagaktiv und besonders in der Jugend in Gruppen lebend. Synodontis hat 
wie die Bodenfische ein unterständiges Maul, ist jedoch pelagisch und nimmt Nah- 


204 ANDREAS HEYD & WOLFGANG PFEIFFER 


rung von der Wasseroberfläche rückenschwimmend auf. S. nigriventris ist dement- 
sprechend sogar invers pigmentiert, wie sein Name verrät. Auch junge Ictaluridae und 
Pimelodidae bilden Schulen. Die pelagischen Eutropiellus, Kryptopterus, Pangasius 
und Plotosus sind tagaktiv und zumindest als Jungtiere ausgesprochene Schwarm- 
fische. Weil die Schreckreaktion ihre Aufgabe nur bei schwarmbildenden, tagaktiven 
Fischen voll erfüllen kann, ist sie bei solitären, nachtaktiven Species nicht zu 
erwarten. Der Schreckstoff dient nicht dem Sender, sondern allein den Empfängern. 
Selektioniert wird nicht das Individuum, sondern die Gruppe. Nur wenn alle 
Schwarmmitglieder gemeinsamer Abstammung wären, kann man von „kin“-Selektion 
sprechen; meist handelt es sich um Gruppen-Selektion. Solitäre, mit kräftigen 
Stacheln bewehrte Fische schützen sich selbst. Im Gegensatz zu den nachtaktiven, 
meist kleinäugigen Welsen sind die tagaktiven Species großäugig, was zu der 
Tatsache paßt, daß die Schreckreaktion visuell übertragen wird. 

Lauterzeugung durch Stridulation ist von mindestens 14 Welsfamilien bekannt, 
Trommelmechanismen von 8 Familien (Tab. 3). In 7 Familien treten beide Mecha- 
nismen auf. Der elektrische Zitterwels besitzt einen Trommelapparat (Müller, 1842), 
ohne zu stridulieren. Er wehrt sich mit seinen starken Entladungen, wie sonst nur 
Zitterrochen und Zitteraal. Von den trommelnden und stridulierenden Familien be- 
sitzen die Mochokidae und Pimelodidae die Schreckreaktion, von den nur stridu- 
lierenden die Ictaluridae und Callichthyidae. Während den stridulierenden Aspredin- 
idae und Loricariidae Schreckstoffzellen und Schreckreaktion fehlen, sind von den 
pelagischen Welsen mit besonders gut ausgebildeter Schreckreaktion nur vereinzelt 
Stridulationsmechanismen bekannt. Schreckreaktion und Lauterzeugung schließen 
einander zwar nicht aus, aber es ist auffallend, daß sie entsprechend der Lebensweise 
der Fische verbreitet sind. Viele nachtaktive, bodenlebende, wehrhafte, solitäre Welse 
können durch Stridulation Abwehrlaute erzeugen. Die tagaktiven, schwarmbildenden 
Welse zeigen eine Schreckreaktion. Doch ist es gegenwärtig wenig ratsam, einen nur 
an wenigen Species erhobenen Befund für eine ganze Familie zu verallgemeinern. 
Aufgrund unvollständiger Untersuchung ist unser bisheriges Bild lückenhaft. Deshalb 
sollten Schlüsse über den Zusammenhang zwischen dem Auftreten von Lauterzeu- 
gungsmechanismen einerseits und dem Vorkommen der Schreckreaktion andererseits 
nur mit Vorsicht gezogen werden. 


DANKSAGUNG 


Wir danken Dr. R. Britz und Dr. H.-J. Franke für Hilfe in taxonomischen 
Fragen sowie Prof. Dr. R. Apfelbach, Prof. Dr. E. Müller, Dr. R. Triebskorn, 
H. Casper und I. Kaipf für technische Unterstützung. 


LITERATUR 


ABU-GIDEIRI, Y. B. & Nasr, D. H. 1973. Sound production by Synodontis schall (Bloch- 
Schneider). Hydrobiologia 43: 415-428. 

Acassız, L. J. R. 1852. (Manner of producing sounds in catfish). 335th Meeting, august 6, 
1850. Proceedings of the American Academy of Arts and Sciences 2: 238. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 205 


AGRAWAL, V. P. & SHARMA, R. S. 1965. Sound producing organs of an Indian catfish, 
Heteropneustes fossilis. The Annals and Magazine of Natural History 13th Series 8: 
339-344. 

ALEXANDER, R. MCN. 1964. The structure of the Weberian Apparatus in the Siluri. Proceedings 
of the Zoological Society of London 142: 419-440. 

ALEXANDER, R. MCN. 1965. Structure and function in the catfish. Journal of Zoology 148: 
88-152. 

ALEXANDER, R. MCN. 1966. Physical aspects of swimbladder function. Biological Reviews 41: 
141-176. 

ARISTOTELES. 1856. Aristoteles Thiergeschichte (pp. 608-609). In: Osiander, C. N. v. & 
Schwab, G. (eds.). Aristoteles Werke. III. Schriften zur Naturphilosophie. Thier- 
geschichte in zehn Büchern, Bd. 2. Verlag der J. B. Metzler ‘schen Buchhandlung, 
Stuttgart. 

Bass, A. H. 1989. Evolution of vertebrate motor systems for acoustic and electric commu- 
nication: peripheral and central elements. Brain, Behavior and Evolution 33: 237-247. 

BERG, L. S. 1958. System der rezenten und fossilen Fischartigen und Fische. VEB Deutscher 
Verlag der Wissenschaften, Berlin, 310 pp. 

BIRKHEAD, W. S. 1972. Toxicity of stings of ariid and ictalurid catfishes. Copeia 1972: 
790-807. 

BOYLE, R. 1670. New pneumatic experiments about respiration. Philosophical Transactions, 
London 5: 2011-2056. 

BOYLE, R. 1675. A conjecture concerning the bladders of air that are found in fishes. 
Philosophical Transactions, London 10: 310-311. 

BREDER, C. M. JR. 1968. Seasonal and diurnal occurences of fish sounds in a small Florida bay. 
Bulletin of the American Museum of Natural History 138: 325-378. 

BRIDGE, T. W. & HADDON, A. C. 1889. Contributions to the anatomy of fishes. I. The air- 
bladder and Weberian ossicles in the Siluridae. Proceedings of the Royal Society of 
London 46: 309-328. 

BRIDGE, T. W. & HADDON, A. C. 1893. Contributions to the anatomy of fishes. II. The air- 
bladder and Weberian ossicles in the siluroid fishes. Philosophical Transactions of the 
Royal Society of London 184: 65-324. 

BRIDGE, T. W. & HADDON, A. C. 1894. Note on the production of sounds by the air-bladder of 
certain siluroid fishes. Proceedings of the Royal Society of London 55: 439-441. 
BRIGGS, J. C. 1979. Ostariophysan zoogeography: an alternative hypothesis. Copeia 1979: 

111-118. 

BROUSSEAU, R. A. 1976. The pectoral anatomy of selected Ostariophysi. Journal of Morpho- 
logy 150: 79-116. 

BURGESS, W. E. 1989. An atlas of freshwater and marine catfishes. A preliminary survey of the 
Siluriformes. 7. F.H. Publications Inc., Neptune City, 784 pp. 

BURKENROAD, M. D. 1931. Notes on the sound producing marine fishes of Louisiana. Copeia 
1931: 20-28. 

CHARDON, M. 1968. Anatomie comparée de l’appareil de Weber et des structures connexes 
chez les Siluriformes. Musée Royal de l’Afrique Centrale-Tervuren, Belgique Annales 
Sciences Zoologiques 169: 1-277. 

CHRANILOV, N. S. 1929. Beiträge zur Kenntnis des Weber’ schen Apparates der Ostariophysi. 2. 
Der Weber’sche Apparat bei Siluroidea. Zoologische Jahrbücher (Anatomie) 51: 
323-462. 

CUVIER, G. & VALENCIENNES, A. 1840. Histoire naturelle des poissons, vol. 15 (p. 187). Verlag 
Pitois-Levrault, Paris. 


206 ANDREAS HEYD & WOLFGANG PFEIFFER 


DAGET, J., GOSSE, J.-P. & THYS VAN DEN AUDENAERDE, D. F. E. 1986. Check-list of the 
freshwater fishes in Africa, vol. 2 (pp. 143-145). ZSNB - Bruxelles, MRAC - Tervuren & 
ORSTOM - Paris, XIV + 520 pp. 

Day, F. 1881. Instincts and emotions in fish. The Journal of the Linnean Society of London 15: 
31-58. 

Day, F. 1958. The fishes of India; being a natural history of the fishes known to inhabit the 
seas and fresh waters of India, Burma and Ceylon, vol. 1 (p. 485). Dawson & Sons, 
London (reprint). 

DEMSKI, L. S., GERALD, J. W. & Poppers, A. N. 1973. Central and peripheral mechanisms of 
teleost sound production. American Zoologist 13: 1141-1167. 

DOBRIN, M. B. 1947. Measurements of underwater noise produced by marine life. Science 105: 
19-23. 

Dorn, E. 1976. Uber den Feinbau von Trommelmuskeln bei Pimelodiden und Doradiden 
(Teleostei, Siluroidei). Verhandlungen der Deutschen Zoologischen Gesellschaft 1976: 
245. 

Duross£, M. 1874. Recherches sur les bruits et les sons expressifs que font entendre les 
poissons d’Europe et sur les organes producteurs de ces phénomenes acoustiques ainsi 
que sur les appareils de l’audition de plusiers de ces animaux. Annales des Sciences 
Naturelles. 5e Série. Zoologie et Paléontologie 20: 115-134. 

EIGENMANN, C. H. & EIGENMANN, R. S. 1890. A revision of the South American Nematognathi 
or cat-fishes. Occasional Papers of the Californian Academy of Sciences 1: 7. 

ESCHMEYER; W. N. (ed.) 1998. Catalog of Fishes. California Academy of Sciences, San Fran- 
cisco, 2905 pp. 

FINE, M. L., FRIEL, J. P., MCELRoy, D., KING, C. B., LOESsER, K. E. & NEWTON, S. 1997. 
Pectoral spine locking and sound production in the channel catfish /ctalurus punctatus. 
Copeia 1997: 777-790. 

FINE, M. L., MCELROY, D., RAFI, J., KING, C. B., LOESSER, K. E. & NEWTON, S. 1996. Latera- 
lization of pectoral stridulation sound production in the channel catfish. Physiology and 
Behavior 60: 753-757. 

FINE, M. L., WINN, H. E. & OLLA, B. L. 1977. Communication in fishes (pp. 472-518). In: 
Sebeok, T. A. (ed.). How animals communicate. /ndiana University Press, Bloo- 
mington, XXI + 1128 pp. 

FINK, S. V. & FINK, W. L. 1981. Interrelationships of the ostariophysan fishes (Teleostei). 
Zoological Journal of the Linnean Society 72: 297-353. 

FINK, S. V. & FINK, W. L. 1996. Interrelationships of ostariophysan fishes (Teleostei) (pp. 209- 
249). In: Stiassny, M. L. J., Parenti, L. R. & Johnson, G. D. (eds.). Interrelationships of 
fishes. Academic Press, New York, XIII + 496 pp. 

FORBES, L. S. 1989. Prey defences and predator handling behaviour: the dangerous prey 
hypothesis. Oikos 55: 155-158. 

FRANKE, H.-J. 1985. Handbuch der Welskunde. Landbuch-Verlag, Hannover, 335 pp. 

FRISCH, K. von. 1923. Ein Zwergwels, der kommt, wenn man ihm pfeift. Biologisches 
Zentralblatt 43: 439-446. 

FRISCH, K. von. 1938. Zur Psychologie des Fisch-Schwarmes. Naturwissenschaften 26: 
601-606. 

FRISCH, K. VON. 1941a. Die Bedeutung des Geruchsinnes im Leben der Fische. Naturwissen- 
schaften 29: 321-333. 

FRISCH, K. von. 1941b. Über einen Schreckstoff der Fischhaut und seine biologische Be- 
deutung. Zeitschrift für vergleichende Physiologie 29: 46-145. 

Frisch, K. von & Stetter, H. 1932. Untersuchungen über den Sitz des Gehörsinnes bei der 
Elritze. Zeitschrift für vergleichende Physiologie 17: 686-801. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 207 


GAINER, H. 1967. Neuromuscular mechanisms of sound production and pectoral spine locking 
in the banjo catfish, Bunocephalus species. Physiological Zoology 40: 296-306. 
GEOFFROY SAINT-HILAIRE, I. 1829. Histoire naturelle des poissons du Nil (pp. 318-319). In: 

Panckouke, C. L. F. (ed.). Description de l’ Egypte, vol. 24. Paris, 579 pp. 

Göhner, M. & Pfeiffer, W. 1996. Über die Verbreitung der Mauthner Axone bei Fischen und 
Amphibien und ihren Zusammenhang mit der Schreckreaktion der Ostariophysi und 
Anura. Revue Suisse de Zoologie 103: 859-891. 

GOSLINE, W. A. 1975. The palatine-maxillary mechanism in catfishes, with comments on the 
evolution and zoogeography of modern siluroids. Occasional Papers of the California 
Academy of Sciences 120: 1-31. 

Greenwood, P. H., Rosen, D. E., Weitzman, S. H. & Myers, G. S. 1966. Phyletic studies of 
teleostean fishes, with a provisional classification of living forms. Bulletin of the 
American Museum of Natural History 131: 339-456. 

Happon, A. C. 1881. On the stridulating apparatus of Callomystax gagata. Journal of Anatomy 
and Physiology 15: 322-326. 

HARDEN JONES, F. R. & MARSHALL, N. B. 1953. The structure and function of the teleostean 
swimbladder. Biological Reviews 28: 16-83. 

HAWKINS, A. D. 1986. Underwater sound and fish behaviour (pp. 114-151). Zn: Pitcher, T. J. 
(ed.). The Behaviour of Teleost Fishes. Croom Helm, London, 553 pp. 

HAWKINS, A. D. & MYRBERG, A. A. 1983. Hearing and sound communication under water 
(pp. 347-405). In: Lewis, B. (ed.). Bioacoustics. A comparative approach. Academic 
Press, London, X + 493 pp. 

INNES, W. T. 1956. Exotic aquarium fishes. /nnes Publishing Company, Philadelphia, ed. 19, 
541 pp. 

KAATZ, I. M. 1995. The evolution of sound signal design in arioid catfishes: swimbladder 
morphology and ecological constraints. American Zoologist 35 (5), 16a. 

KAATZ, I. M. 1999. The behavioral and morphological diversity of sound communication 
systems in a clade of tropical catfishes, with comparison to ten additional acoustic 
catfish families (Order Siluriformes, superfamily Arioidei). Doctoral dissertation, Syra- 
cuse, New York, 365 pp. 

KAATZ, I. M. & LOBEL, P. S. 1999. Acoustic behavior and reproduction in five species of 
Corydoras catfishes (Callichthyidae). Biological Bulletin, 197: 241. 

KAATZ, I. M. & STEWART, D. J. 1997. The evolutionary origin and functional divergence of 
sound production in catfishes: stridulation mechanisms. American Zoologist 37, 137a. 

KARLSON, P. M. & LÜSCHER, M. 1959. „Pheromones“: a new term for a class of biologically 
active substances. Nature (London) 183: 55-56. 

KASTBERGER, G. 1977. Der Trommelapparat der Doradidae (Siluriformes, Pisces). Zoologische 
Jahrbücher: Allgemeine Zoologie und Physiologie 81: 281-309. 

KASTBERGER, G. 1978. Producäo de sons em Doradideos e Auchenopterideos (Siluriformes, 
Pisces). Acta Amazonica 8: 455-468. 

KRATOCHVIL, H., SCHACHNER, G. & VÖLLENKLE, W. 1980. Stridulations- und Trommelmus- 
kellaute beim Aggressionsverhalten von Pimelodus sp. (Siluridae, Pisces). Wissen- 
schaftlicher Film Wien 25: 23-27 (Ctf 1634). 

KRATOCHVIL, H. & VÖLLENKLE, W. 1981. Bewegung der Brustflossen beim Stridulieren von 
Pimelodus sp. (Siluridae, Pisces). Wissenschaftlicher Film Wien 26: 49-53 (Z 1627). 

LADICH, F. 1997. Comparative analysis of swimbladder (drumming) and pectoral (stridulation) 
sounds in three families of catfishes. Bioacoustics 8: 185-208. 

LADICH, F. & Fine, M. L. 1994. Localization of swimbladder and pectoral motoneurons in- 


volved in sound production in pimelodid catfish. Brain, Behavior and Evolution 44: 
86-100. 


208 ANDREAS HEYD & WOLFGANG PFEIFFER 


LAUDER, G. V. & LIEM, K. F. 1983. The evolution and interrelationships of the actinopterygian 
fishes. Bulletin of the Museum of Comparative Zoology 150: 95-197. 

LILEY, N. R. 1982. Chemical communication in fish. Canadian Journal of Fisheries and 
Aquatic Sciences 39: 22-35. 

LUNDBERG, J. G. 1992. The phylogeny of Ictalurid catfishes: a synthesis of recent work 
(pp. 392-420). In: Mayden, R. L. (ed.). Systematics, historical ecology, and North 
American freshwater fishes. Stanford University Press, Stanford, XX VI + 969 pp. 

LUNDBERG, J. G. & MCDADE, L. A. 1986. On the south american catfish Brachyrhamdia imi- 
tator Myers (Siluriformes, Pimelodidae), with phylogenetic evidence for a large intra- 
familial lineage. Notulae naturae of the Academy of Natural Sciences of Philadelphia 
463: 1-24. 

MAHAJAN, C. L. 1963. Sound producing apparatus in an Indian catfish Sisor rhabdophorus 
Hamilton. Journal of the Linnean Society of London 44: 721-724. 

MARKL, H. 1968. Das Schutzverhalten eines Welses (Hassar orestis Steindachner) gegen 
Angriffe von Piranhas (Serrasalmus nattereri Kner). Zeitschrift fiir Tierpsychologie 26: 
385-389. 

Mayr, M. 1987. Verhaltensbeobachtungen an Hoplosternum thoracatum, mit besonderer 
Berücksichtigung des Lautverhaltens. Dissertation, Wien, 103 pp. 

MICHELSEN, A. 1983. Biophysical basis of sound communication (pp. 3-38). In: Lewis, B. 
(ed.). Bioacoustics. A comparative approach. Academic Press, London, X + 493 pp. 

Mo, T. 1991. Anatomy, relationships and systematics of Bagridae (Teleostei) with a hypothesis 
of siluroid phylogeny. Koeltz Scientific Books, Koenigstein, VII + 216 pp. 

MONTOYA-BURGOS, J.-L, MULLER, S., WEBER, C. & PAWLOWSKI, J. 1998. Phylogenetic rela- 
tionships of the Loricariidae (Siluriformes) based on mitochondrial rRNA gene 
sequences (pp. 363-374). In: Malabarba, L. R., Reis, R. E., Vari, R. P., Lucena, Z., M. 
S. & Lucena, C. A. S. (eds.). Phylogeny and classification of neotropical fishes. 
EDIPUCRS, Porto Alegre, X + 603 pp. 

Moreau, A. 1876. Recherches expérimentales sur les fonctions de la vessie natatoire. Annales 

des Sciences naturelles; 6. Série, Zoologie et Paléontologie 4: 1-85. 

MULLER, J. 1842. Beobachtungen tiber die Schwimmblase der Fische, mit Bezug auf einige 

neue Fischgattungen. Archiv fiir Anatomie, Physiologie und wissenschaftliche Medizin 

1842: 307-329. 

MÜLLER, J. 1857. Uber die Fische, welche Töne von sich geben und die Entstehung dieser 

Töne. Archiv für Anatomie, Physiologie und wissenschaftliche Medizin 1857: 249-279. 

MYRBERG, A. A. 1981. Sound communication and interception in fishes (pp. 395-426). In: 
Tavolga, W. N., Popper, A. N. & Fay, R. R. (eds.). Hearing and Sound Communication 
in Fishes. Springer, New York, XVI + 608 pp. 

NAWAR, G. 1954. On the anatomy of Clarias lazera. 1. Osteology. Journal of Morphology 94: 
551-585. 

NELSON, J. S. 1994. Fishes of the world. John Wiley, New York, ed. 3, 416 pp. 

Novacek, M. J. & MARSHALL, L. G. 1976. Early biogeographic history of ostariophysan fishes. 
Copeia 1976: 1-12. 

PFEIFFER, W. 1960. Uber die Schreckreaktion bei Fischen und die Herkunft des Schreckstoffes. 
Zeitschrift fiir vergleichende Physiologie 43: 578-614. 

PFEIFFER, W. 1962a. Uber die Verbreitung der Schreckreaktion bei Salmlern (Characidae) und 
Welsen (Siluroidea). Naturwissenschaften 49: 614. 

PFEIFFER, W. 1962b. The fright reaction of fish. Biological Reviews 37: 495-511. 

PFEIFFER, W. 1963a. Vergleichende Untersuchungen über die Schreckreaktion und den 
Schreckstoff der Ostariophysen. Zeitschrift fiir vergleichende Physiologie 47: 111-147. 

PFEIFFER, W. 1963b. Alarm substances. Experientia 19: 113-123. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 209 


PFEIFFER, W. 1966. Die Schreckreaktion der Fische und Kaulquappen. Naturwissenschaften 22: 
565-570. 

PFEIFFER, W. 1967. Schreckreaktion und Schreckstoffzellen bei Ostariophysi und Gonorhynchi- 
formes. Zeitschrift fiir vergleichende Physiologie 56: 380-396. 

PFEIFFER, W. 1970. Uber die Schreckstoffzellen der Siluriformes. Anatomischer Anzeiger 126: 
113-119. 

PFEIFFER, W. 1974. Pheromones in fish and amphibia (pp. 269-296). In: Birch, M. C. (ed.). 
Pheromones. North-Holland Publishing Company, Amsterdam, XXI + 495 pp. 

PFEIFFER, W. 1977. The distribution of fright reaction and alarm substance cells in fishes. 
Copeia 1977: 653-665. 

PFEIFFER, W. 1982. Chemical signals in communication (pp. 307-326). In: Hara, T. J. (ed.). 
Chemoreception in fishes. Elsevier, Amsterdam, X + 433 pp. 

PFEIFFER, W., DENOIX, M., WEHR, R., GNASS, D., ZACHERT, I. & BREISCH, M. 1986. Video- 
technische Verhaltensanalyse der Schreckreaktion von Ostariophysen (Pisces) und die 
Bedeutung des „Mauthner-Reflexes“. Zoologische Jahrbücher, Allgemeine Zoologie 
und Physiologie der Tiere 90: 115-165. 

PFEIFFER, W. & EISENBERG, J. F. 1965. Die Lauterzeugung der Dornwelse (Doradidae) und der 
Fiederbartwelse (Mochokidae). Zeitschrift für Morphologie und Okologie der Tiere 54: 
669-679. 

PINNA DE, M. C. C., 1998. Phylogenetic relationships of neotropical Siluriformes (Teleostei: 
Ostariophysi): Historical overview and synthesis of hypotheses (pp. 279-330). In: Mala- 
barba, L. R., Reis, R. E., Vari, R. P., Lucena, Z. M. S. & Lucena, C. A. S. (eds.). Phylo- 
geny and classification of neotropical fishes. EDIPUCRS, Porto Alegre, X + 603 pp. 

POGGENDORF, D. 1952. Die absoluten Hörschwellen des Zwergwelses (Amiurus nebulosus) und 
Beiträge zur Physik des Weberschen Apparates der Ostariophysen. Zeitschrift für ver- 
gleichende Physiologie 34: 222-257. 

Popper, A. N. & FAY, R. R. 1973. Sound detection and processing by teleost fishes: a critical 
review. Journal of the Acoustical Society of America 53: 1515-1529. 

Popper, A. N. & Fay, R. R. 1993. Sound detection and processing by fish: critical review and 
major research questions. Brain, Behavior and Evolution 41: 14-38. 

Popper, A. N. & TAVOLGA, W. N. 1981. Structure and function of the ear in the marine catfish, 
Arius felis. Journal of Comparative Physiology 144: 27-34. 

PRUZSINSZKY, I. & LADICH, F. 1998. Sound production and reproductive behaviour of the 
armoured catfish Corydoras paleatus (Callichthyidae). Environmental Biology of 
Hishes 53: 183-191. 

Ray, J. 1675. A letter.....about the swiming bladders in fishes. Philosophical Transactions, 
London 10: 349-351. 

REGAN, C. T. 1911. The classification of the teleostean fishes of the order Ostariophysi. - 2. 
Siluroidea. Annals and Magazine of Natural History 8: 553-577. 

REISSNER, E. 1859. Uber die Schwimmblase und den Gehörapparat einiger Siluroiden. Archiv 
für Anatomie, Physiologie und wissenschaftliche Medizin 1859: 421-438. 

RIEHL, R. & BAENSCH, H. A. 1983. Aquarienatlas. Bd. 1. Mergus Verlag, Melle, 992 pp. 

RIGLEY, L. & Muir, J. 1979. The role of sound production by the brown bullhead, Jctalurus 
nebulosus. Proceedings of the Pennsylvania Academy of Science 53: 132-134. 

ROBERTS, T. R. 1973. Interrelationships of ostariophysans (pp. 373-395). In: Greenwood, P. H., 
Miles, R. S. & Patterson, C. (eds.). Interrelationships of fishes. Academic Press, 
London, XVI + 536 pp. 

Rosen, D. E. & GREENWOOD, P. H. 1970. Origin of the Weberian Apparatus and the rela- 
tionships of the ostariophysan and gonorynchiform fishes. American Museum Novitates 
2428: 1-25. 


210 ANDREAS HEYD & WOLFGANG PFEIFFER 


SAGEMEHL, M. 1885. Beiträge zur vergleichenden Anatomie der Fische. III. Das Cranium der 
Characiniden nebst allgemeinen Bemerkungen über die mit einem Weber’schen Appa- 
rat versehenen Physostomenfamilien. Morphologisches Jahrbuch 10: 1-119. 


SCHACHNER, G. 1977. Mechanismen und biologische Bedeutung der Schallerzeugung und 
- wahrnehmung beim südamerikanischen Antennenwels (Pimelodus sp., Pimelodidae). 
Dissertation, Wien, 103 pp. 


SCHACHNER, G. & SCHALLER, F. 1982. Schallerzeugung und Schallreaktionen beim Antennen- 
wels (Mandim) Rhamdia sebae sebae. Zoologische Beiträge 27: 375-391. 


SCHAEFER, S. A. 1984. Mechanical strength of the pectoral spine / girdle complex in Pterygopl- 
ichthys (Loricariidae: Siluroidei). Copeia 1984: 1005-1008. 

SCHAEFER, S. A. 1987. Osteology of Hypostomus plecostomus (L.), with a phylogenetic 
analysis of the Loricariid subfamilies (Pisces: Siluroidei). Contributions in Science. 
Natural History Museum of Los Angeles County 394: 1-31. 


SCHELLART, A. M. & WUBBELS, R. J. 1998. The auditory and mechanosensory lateral line 
system, pp. 283-312. In: Evans, D. H. (ed.). The physiology of fishes. CRC Press, New 
York, 519 pp. 

SCHNEIDER, H. 1961. Neuere Ergebnisse der Lautforschung bei Fischen. Naturwissenschaften 
48: 513-518. 

SCHNEIDER, H. 1967. Morphology and physiology of sound-producing mechanisms in teleost- 
fishes. Marine Bio-Acoustics 2: 135-158. 

SCHUTZ, F. 1956. Vergleichende Untersuchungen über die Schreckreaktion bei Fischen und 
deren Verbreitung. Zeitschrift für vergleichende Physiologie 38: 84-135. 

SMITH, R. J. F. 1973. Testosterone eliminates alarm substance in male fathead minnows. Cana- 
dian Journal of Zoology 51: 875-876. 

SMITH, R. J. F. 1976a. Seasonal loss of alarm substance cells in North American cyprinoid 
fishes and ist relation to abrasive behaviour. Canadian Journal of Zoology 54: 1172- 
1182. 

SMITH, R. J. F. 1976b. Male fathead minnows (Pimephales promelas Rafinesque) retain their 
fright reaction to alarm substance during the breeding season. Canadian Journal of 
Zoology 54: 2230-2231. 

SMITH, R. J. F. 1977. Chemical communication as adaptation: alarm substance in fish (pp. 303- 
320). In: Müller-Schwarze, D. & Mozell, M. M. (eds.). Chemical signals in vertebrates. 
Plenum Press, New York, X + 609 pp. 

SMITH, R. J. F. 1982. The adaptive significance of the alarm substance - fright reaction system 
(pp. 327-342). In: Hara, T. J. (ed.). Chemoreception in fishes. Elsevier, Amsterdam, X + 
433 pp. 

SMITH, R. J. F. 1986. Reduced alarm substance cell counts in male and androgen-treated zebra 
danios (Brachydanio rerio). Canadian Journal of Zoology 64: 551-553. 

SMITH, R. J. F. 1992. Alarm signals in fishes. Reviews in Fish Biology and Fisheries 2: 33-63. 

SORENSEN, W. 1884. Om Lydorganer hos Fiske. En physiologisk og comparativ-anatomisk 
undersögelse. V. Thaning & Appels Boghandel, Copenhagen, 245 pp. 

SÖRENSEN, W. 1895. Are the extrinsic muscles of the air-bladder in some Siluroidae and the 
„elastic-spring“ apparatus of others subordinate to the voluntary production of sounds? 
What is, according to our present knowledge, the function of the Weberian ossicles? A 
contribution to the biology of fishes. Journal of Anatomy and Physiology 29: 109-139, 
205-229, 399-423, 518-552. 

SORENSEN, W. 1898. Some remarks on Dr. Thilo’s memoir on „Die Umbildungen an den 
Gliedmaßen der Fische“. Morphologisches Jahrbuch 25: 170-189. 

SOLOMON, D. J. 1977. A review of chemical communication in freshwater fish. Journal of Fish 
Biology 11: 363-376. 


LAUTERZEUGUNG DER WELSE (SILUROIDEI) 211 


STABELL, O. B. 1996. Alarm signals as primer pheromones in fishes (pp. 39-46). In: Canario, 
A. V. M. & Power, D. M. (compilers). Fish Pheromones. Proceedings of a Workshop at 
University of Algarve, Faro, Portugal, 22-24 May 1995. 

STERBA, G. 1956. Süßwasserfische aus aller Welt. Verlag Zimmer und Herzog, Berchtesgaden, 
638 pp. 

STERBA, G. 1990. Süßwasserfische der Welt. Urania-Verlag, Leipzig, 915 pp. 

STEWART, D. J. 1986. Revision of Pimelodina and description of a new genus and species from 
the Peruvian Amazon (Pisces: Pimelodidae). Copeia 1986: 653-672. 

TAVOLGA, W. N. 1960. Sound production and underwater communication in fishes (pp. 93- 
136). In: Lanyon, W. E. & Tavolga, W. N. (eds.). Animal sounds and communication. 
American Institute of Biological Sciences, XIII + 443 pp. 

TAVOLGA, W. N. 1962. Mechanisms of sound production in the Ariid catfishes Galeichthys and 
Bagre. Bulletin of the American Museum of Natural History 124: 1-30. 

TAVOLGA, W. N. 1971a. Sound production and detection (pp. 135-205). In: Hoar, W. S. & 
Randall, D. J. (eds.). Fish Physiology, vol. 5. Academic Press, New York, XVI + 
600 pp. 

TAVOLGA, W. N. 1971b. Acoustic orientation in the sea catfish Galeichthys felis. Annals of the 
New York Academy of Sciences 188: 80-97. 

TAVOLGA, W. N. 1976. Acoustic obstacle detection in the sea catfish (Arius felis) (pp. 185-204). 
In: Schuijf, A. & Hawkins, A. D. (eds.). Sound reception in fish. Elsevier, Amsterdam, 
VII + 287 pp. 

TAVOLGA, W. N. 1977. Sound production and detection (pp. 3-53). In: Tavolga, W. N. (ed.). 
Sound production in fishes. Benchmark Papers in Animal Behavior, vol. 9; Dowden, 
Hutchinson & Ross, Stroudsburg, XII + 363 pp. 

TENNENT, J. E. 1859. Ceylon: an account of the island, physical, historical, and topographical 
with notices of its natural history, antiquities and productions (p. 470). Longman, 
Green, Longman & Roberts London, 2nd edition, Vol. 2. 

TEUGELS, G. G. 1996. Taxonomy, phylogeny and biogeography of catfishes (Ostariophysi, 
Siluroidei): an overview. Aquatic living resources, vol. 9, Hors série: 9-34. 

Tyack, P. L. 1998. Acoustic communication under the sea (pp. 163-220). In: Hopp, S. L, 
Owren, M. J. & Evans, C. S. (eds.). Animal acoustic communication. Springer, Heidel- 
berg, XIX + 421 pp. 

URICK, R. J. 1983. Principles of underwater sound. McGraw-Hill, New York, XHI + 423 pp. 

VILLWOCK, W. 1960. Zur Lauterzeugung bei Knochenfischen, unter besonderer Berücksich- 
tigung von Acanthodoras spinosissimus. Deutsche Aquarien und Terrarien Zeitschrift 
13: 237-240. 

WEBER, E. H. 1820. De aure et auditu hominis et animalium. I. De aure animalium aquatilium. 
Lipsiae, 134 pp. 

WEISS, B. A., STROTHER, W. F. & HARTIG, G. M. 1969. Auditory sensitivity in the bullhead 
catfish Uctalurus nebulosus). Proceedings of the Natural Academy of Sciences of the 
U.S.A. 64: 552-556. 

WINN, H. E. 1964. The biological significance of fish sounds (pp. 213-231). In: Tavolga, W.N. 
(ed.). Marine bio-acoustics. Pergamon Press, Oxford, XII + 413 pp. 

WRIGHT, R. R. 1884. The relationship between the air-bladder and auditory organ in Amiurus. 
Zoologischer Anzeiger 7: 248-252. 

ZELICK, R., MANN, D. A. & Popper, A. N. 1999. Acoustic communication in fishes and frogs 
(pp.363-411). In: Fay, R. R. & Popper, A. N. (eds.). Comparative hearing: fish and 
amphibians. Springer, New York, XVIII + 438 pp. 


ws 


am ITA eae a ES 


ol i bu 
n. Tr VE vei (ES: 
n | LPS VE UNITI wik 


el TE ME ihe} 
“ALE sth: ie 


Ji TAN 
re 


FE x 

rs | hi 

ar M 
rite AI tr 5 iv IK à 


à Aut das 
ripa wd 27 | 


REVUE SUISSE DE ZOOLOGIE, 107 (1): 213-232; mars 2000 


Molecular and morphological evidence for the presence of a 
new Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus 


Benjamin GANTENBEIN*!, Christian KROPF?, Carlo Rodolfo LARGIADER! 
& Adolf SCHOLL! 
| Division of Population Biology, Institute of Zoology, University of Berne, 
Baltzerstrasse 3, CH-3012, Berne, Switzerland. 
e-mail: Gantenbein @zoo.unibe.ch 
2 Natural History Museum Berne, Department of Invertebrates, 
Bernastrasse 15, CH-3005 Berne, Switzerland. 
e-mail: christian.kropf@nmbe.unibe.ch 


Molecular and morphological evidence for the presence of a new 
Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus. - Allo- 
zyme data (16 loci) from Mesobuthus gibbosus populations in the eastern 
Mediterranean region show that the Cyprus population is highly distinct, 
although morphological differentiation is rather weak. This provides 
evidence for a ‘hidden’ taxon on Cyprus. This island population 1s described 
as a new species. 


Key-words: Allozymes - Scorpiones - Mesobuthus — new species - Cyprus. 


INTRODUCTION 


Comparison of molecular data from insular and continental populations of a 
species or of closely related species may provide new insights into the process of 
speciation with respect to time (Estoup er al., 1996; Baldwin & Sanderson, 1998; Beerli 
et al., 1996; Gillespie et al., 1998; Hollocher, 1998; Widmer er al., 1998). Island 
populations are of particular interest for the analysis of colonisation processes (Vachon 
& Abe, 1988) or for studying founder effect speciation (Templeton, 1980, 1981; Carson 
& Templeton, 1984; Grant, 1998). Mesobuthus gibbosus (Brullé, 1832) (Scorpiones: 
Buthidae) is widely distributed in the eastern Mediterranean region (Werner, 1928, 
1937, 1938; Vachon, 1947a, 1947b, 1948, 1966; Tolunay, 1959; Gruber, 1963, 1966; 
Kinzelbach, 1975, 1982, 1984, 1985; Vachon & Kinzelbach, 1987; Kritscher, 1993; 
Crucitti, 1993). Its geographical range includes the mainland (south Balkan, Pelo- 
ponnesus, Anatolia) and eastern Mediterranean islands (e. g. Cyclades, Sporades, Crete 


“to whom correspondence should be addressed 
Manuscript accepted 05.10.1999 


214 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


and Cyprus). The geographic variation within scorpion species was traditionally exami- 
ned by using morphological characters like patterns of trichobothria (=‘tricho- 
bothriotaxy’) (Vachon, 1974, 1975, 1976; Fet, 1986; Fet & Rechkin, 1989) or variation 
in the number of pectinal teeth (Kinzelbach, 1975; Michalis & Kattoulas, 1981; 
Michalis & Dolkeras, 1989). The taxonomic status of different populations of M. 
gibbosus has remained unclear: Kinzelbach (1975) distinguished two subspecies, M. g. 
gibbosus (Brullé, 1832) (south of the Balkan Peninsula, Northern Sporades, Cyclades) 
and M. g. anatolicus (Schenkel, 1947) (Crete, Anatolia, Cyprus). Kritscher (1993) 
doubted the validity of M. g. anatolicus and rejected it. 

The widely accepted theory of the salinity crisis (Hsü, 1972; Hsü et al., 1977) 
states that the Mediterranean sea dried out 5.6 Myrs ago and, consequently, the 
colonisation of islands via landbridges was possible during a period of 100°000 yrs. 
Since the refilling of the basin (5.2 Myrs ago) the populations became geographically 
isolated. Since then, a considerable degree of differentiation between mainland and 
island populations is expected to have taken place. Alternatively, other genetic 
population structures than expected are likely if scorpions were introduced by man. 
Because of several introductions of euscorpiids (Stockwell 1992) caused by man (e. g. 
Benton, 1991; Goyffon, 1992; Toscano-Gadea, 1998), island populations might not be 
as isolated as expected. 

To clarify the status of some island populations and of the Cyprus population in 
particular, we carried out a comparative genetic analysis at 16 allozyme loci. 
Populations from the Peloponnesus, from the south of the Balkans, from Crete, Rhodes, 
Anatolia and from Cyprus were included. We describe here new morphological 
characters, which distinguish Mesobuthus from Cyprus from the other populations of 
Mesobuthus gibbosus examined. 


MATERIAL & METHODS 


Specimens analysed. Samples were collected at two sites on the Peloponnesus, 
GR (Vigla [Arta]; Mathia [Messinia]), two sites on Crete, GR (Vai; Zakros [both in 
Lassithi]), two sites on Rhodes, GR (Petaloudes: Kolympia), two sites in central 
Anatolia, TR (Avanos; Hacibectas [both Cappadocia]), one site in southern Anatolia, 
TR (Selale [Pamphylia]) and two sites on northern Cyprus (Tepebasi; Kantara [both 
Turkish part]). The scorpions were transported alive to the laboratory, killed by deep- 
freezing and stored at -80°C prior to electrophoresis. For the morphological exami- 
nation, the specimens were later transferred into 75% ethanol. Androctonus maure- 
tanicus (Pocock, 1902) from Agadir (MA) and Androctonus crassicauda (Olivier, 
1807) from Urfa (TR) were used as outgroup species for phylogenetic analyses. The 
sampling sites are shown in Fig. 1. 

Allozyme analysis. Horizontal starch gel electrophoresis of allozymes was 
carried out according to the protocols described in Gantenbein er al. (1998) and Murphy 
et al. (1996). We scored 16 loci on three buffer systems: N-(3-Aminopropyl)-morpho- 
line-citrate (AC, pH 6.2, modified from Clayton & Tretiak, 1972), Tris-citrate (TC, pH 
7.3, Ayala et al., 1972) and Tris-borate-EDTA (TBE, pH 9.3, modified from Ayala er 


A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 


(N°) 
Nn 


Zo & 
The BALKANS 


= 


\ 
ne 


SS) 5 


4 


FIG. | 


Sampling sites of the analysed Mesobuthus populations: 1: Vigla, GR, 2: Mathia, Gr, 3: Vai, GR, 
4: Zakros, GR, 5: Kolympia, GR, 6: Petaloudes, GR, 7: Selale, TR (3 sites), 8: Hacibectas, TR, 9: 
Avanos, TR, 10: Kantara, TR, 11: Tepebasi, TR. 


al., 1972). The loci scored were: ALPDH (Alanopine dehydrogenase; EC 1.5.1.17), 
ARK (arginine kinase; EC 2.7.3.3), AAT-1 and AAT-2 (aspartate aminotransferase; EC 
2.6.1.1), DDH (dihydrolipoamide oxidase; EC 1.8.1.4), PGI (GPI) (glucose-6-phos- 
phate isomerase; EC 5.3.1.9), GTDH (glutamate dehydrogenase; EC 1.4.1.2), IDH-1 
and IDH-2 (isocitrate dehydrogenase; EC 1.1.1.42), MDH-I and MDH-2 (malate 
dehydrogenase; EC 1.1.1.37), MPI (mannose-6-phophate isomerase; EC 5.3.1.8), PGM 
(phosphoglucomutase; EC 5.4.2.2), 6-PGD (6-phosphogluconate dehydrogenase; EC 
1.1.1.44), PK (pyruvate kinase; EC 2.7.1.40), and SOD (superoxide dismutase; EC 
1.15.1.1). We refer to the observed electromorphs as alleles which are identified by 
their electrophoretic mobility relative to the most common mobility in the Euscorpius 
flavicaudis (de Geer, 1778) population from Lauris, France (assigned mobility=100) as 
described in Gantenbein er al. (1998). To assess the genetic variability within each 
population, the mean number of alleles per locus, the percentage of polymorphic loci 
and the mean heterozygosity were calculated by the direct count method and by Nei’s 
(1978) unbiased estimate. Nei’s genetic distance (1972) was calculated from pairwise 


216 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


comparisons of populations using the program GENDIST from the PHYLIP 3.5 
package (Felsenstein, 1995). Nei’s distance is expected to rise linearly with time since 
the complete separation of gene pools, if sufficiently large numbers of loci are 
measured (Felsenstein 1984; Nei 1987). Using Nei’s pairwise distances as an input 
matrix, an additive tree was created by the neighbour-joining algorithm (NJ) (Saitou & 
Nei, 1987). NJ is assumed to be a good heuristic approach for estimating the minimum 
evolution tree (Page & Holmes, 1998) and allows for different rates among lineages in 
contrast to the UPGMA clustering algorithm. Alternatively, an unrooted maximum 
likelihood tree was calculated using the computer program CONTML. It estimates 
phylogenies by the restricted maximum likelihood (REML) method, based on the 
Brownian motion model (Cavalli-Sforza & Edwards, 1967). The REML algorithm was 
formerly described in Felsenstein (1973, 1981) and uses less parameters than the full 
ML analysis and is, therefore, assumed to be more consistent. Additionally, the 
program calculates branch lengths and rough confidence intervals for the branches. 
Bootstrap values were obtained based on 1000 pseudo-replicates of allele frequencies 
using the SEQBOOT routine in PHYLIP. 


RESULTS 


Genetic variability. In general, the observed genetic variability was rather low 
within the Mesobuthus samples. The mean observed heterozygosity was 0.07 + 0.05, 
one to six out of 16 loci were polymorphic at the 0.95 criterion. The mean number of 
alleles ranged from 1.0 to 1.5 per locus. All analysed populations were fixed at many 
loci (Table 1). However, this variation was not evenly distributed among Mesobuthus 
samples. At five loci, more than two electromorphs were detected, whereas all 
populations were fixed for the same allele at three loci (Idh-2, Mdh-1, Pk). The Cyprus 
samples were fixed for private alleles (alleles that were not found in any other 
population) or showed polymorphisms with private alleles at six loci (Aat-1, Aat-2, 
Alpdh, Ldh-2, Mdh-2, 6-Pgd) in addition to minor differences in allele frequencies at 
the locus Mpi (Table 1). On the other hand, if compared to the mainland populations, 
the island populations from Crete and Rhodes showed private alleles at one locus only 
(samples Zakros and Vai at Aat-1 and samples Kolympia and Petaloudes at 6-Pgd, 
respectively). The island samples and the mainland samples differed mainly in the 
allele frequencies. 

The outgroup comparison with A. mauretanicus and A. crassicauda, respecti- 
vely, revealed alleles at eight loci that were not found in Mesobuthus, whereas, with 
respect to allele frequencies, the two Androctonus species differed from each other only 
at two loci. 


TABLE | 


Allele frequencies and sample sizes of eleven Mesobuthus populations. Androctonus maure- 
tanicus (MA) and A. crassicauda (TR) were used as outgroup species. Alleles were labelled as 
described in the material & methods section. Measures of genetic variability for each population 
are given at the bottom of the table. Private alleles of the Cyprus samples and the heterozygosity 


Genus/species 
Region 


Site 


Sample size 


Locus Allele 


Aat-1 111 


Aat-2 117 


Alpdh 105 
Ark 102 
Ddh 99 
Gtdh 100 


Idh-1 104 


Idh-2 93 
Mdh-1 105 


Mdh-2 92 


Mpi 160 


6-Pgd 112 


Pgi (GPI) 95 
87 
Pgm 88 
81 


Pk 100 
98 


Sod 110 
109 
103 

Mean no. of 

alleles per locus 


Percentage of 


polymorhphic loci 


Heterozygosity 
(observed) 
Heterozygosity 
(expected) 


Greek mainland 


A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 


M. gibbosus 


Crete 


Vigla Mathia Zakros Vai 


(2) 


1.00 


1.00 
0.03 


1.00 


1.00 


1.00 


1.00 


1.00 


1.00 


1.00 


1.00 


0.25 
0.25 


0.50 


1.00 


0.25 
0.75 


1.00 


1.00 


0.06 
(0.0) 


0.08 
(0.1) 


(15) 


1.00 


0.97 


1.00 


1.00 


1.00 


1.00 


1.00 


1.00 


1.00 


1.00 
1.00 


1.00 


(5) 


1.00 


1.00 
1.00 
1.00 


1.00 
1.00 


1.00 


1.00 


1.00 


1.00 


0.50 
0.50 


1.00 


1.00 
1.00 


1.00 
1.00 


1.1 
(0.0) 


6.3 


0.01 
(0.0) 


0.04 
(0.0) 


(2) 


1.00 


1.00 


1.00 


1.00 


1.00 
1.00 


1.00 


1.00 


1.00 


1.00 


1.00 


1.00 


Rhodes 


bia 
(6) 


1.00 


1.00 


1.00 


0.67 
0.33 


1.00 
0.67 


0.33 


1.00 


1.00 


1.00 


1.00 
1.00 


0.08 


0.92 


Kolym- Peta- 
loudes 


(6) 


1.00 


1.00 
0.09 


1.00 


22 


oo — 
wn 


1.00 
0.25 
0.75 
0.08 
0.92 


1.00 


1.00 


1.00 
1.00 


S. Ana- 
tolia 


Selale 


(16) 


1.00 


0.91 


1.00 


1.00 


1.00 


1.00 


0.81 


0.19 
1.00 


1.00 


1.00 


0.97 
0.03 


1.00 


C. Ana- 
tolia 
Avanos Haci- 
bectas 
(5) (3) 
1.00 1.00 
1.00 1.00 
1.00 1.00 
1.00 1.00 
1.00 1.00 
1.00 1.00 
0.90 1.00 
0.10 
1.00 1.00 
1.00 1.00 
1.00 1.00 
0.90 1.00 
0.10 
1.00 1.00 
1.00 1.00 
1.00 1.00 
1.00 1.00 
1.00 1.00 
1.1 1.0 
(0.1) (0.0) 
12.5 0.0 
0.03 0.0 
(0.0) (0.0) 
0.03 0.0 
(0.0) (0.0) 


M. cyprius 
Cyprus 
Tepe- Kan- 
basi tara 
(20) (4) 
0.30 
0.70 1.00 
0.35 0.63 
0.13 
0.65 0.24 
1.00 1.00 
1.00 1.00 
0.05 
0.95 1.00 
1.00 1.00 
0:977721:00 
0.03 
1.00 1.00 
1.00 1.00 
1.00 1.00 
0.62 1.00 
0.38 
0.03 
0.12 
0.94 0.88 
0.03 
0.08 
0.92 1.00 
1.00 1.00 
1.00 1.00 
1.00 1.00 
1:5 12 
(0.2) (0.1) 
Is NAS 
0.08 0.06 
(0.0) (0.0) 
0.11 0.05 
(0.1) (0.0) 


Aourir 


(14) 


1.00 


1.00 


1.00 
1.00 


1.00 


1.00 


1.00 


1.00 
1.00 


0.04 
0.96 


1.00 


1.00 


0.03 
(0.0) 


0.04 
(0.0) 


217 


Androctonus 
Morocco Turkey 


Urfa 
(6) 


0.08 
0.75 
0.17 


1.00 


1.00 
1.00 


1.00 


1.00 


1.00 


1.00 
1.00 


0.67 
0.33 


1.00 


1.00 


0.09 
(0.1) 


0.09 
(0.1) 


218 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


TABLE 2 


Calculated pairwise distance matrix (Nei’s D 1972) for all 13 samples of Mesobuthus. In bold are 
the genetic distance values of pairwise comparisons that involved one of the two Cyprus samples. 


Region Greek mainland Crete Rhodes Anatolia Cyprus Outgroup 

Population Vigla Mathia Zakros Vai Kolympia Peta- Selale Avanos Haci- Tepe- Kantara Aourir 
loudes bectas basi 

Vigla 

Mathia 0.03 

Zakros 0.42 0.41 

Vai 0.44 0.47 0.02 

Kolympia 0.13 0.10 0.21 0.26 

Petaloudes 0.05 0.03 0.35 0.42 0.05 

Selale 0.45 0.39 0.16 0.21 0.19 0.33 

Avanos 0.34 0.29 0.23 0.28 0.11 0.24 0.07 

Hacibectas 0.34 0.29 0.23 0.29 0.10 0.23 0.07 0.00 

Tepebasi 0.31 0.28 0.78 0.84 0.44 0.33 0.87 0.70 0.69 

Kantara 0.39 0.33 0.82 0.92 0.50 0.39 0.95 0.78 0.77 0.03 

Aourir 1.40 [ESI 1.13 IIS 1.09 1.32 0.95 0.80 0.81 1.32 135 

Urfa 1.49 1.55 1.26 1.28 1.11 1.30 1.04 0.88 0.90 1.44. 1.46 0.14 


Genetic differentiation and phylogenetic analyses. The calculated Nei’s dis- 
tances from pairwise comparisons of populations within Mesobuthus ranged from 0.00 
to 0.95 (Table 2). The distance values between the two outgroup species and Meso- 
buthus were at least 0.80 and ranged up to 1.55. If the two samples from Cyprus are 
compared with the other Mesobuthus samples, it is obvious that the Cyprus populations 
are considerably distinct from all others, 1. e. Vigla, Mathia, Kolympia, Petaloudes, 
Zakros, Vai (all GR), Selale, Avanos, Hacibectas (all TR), by rather high distance 
values (0.31 - 0.95). 

The topologies of the NJ tree and of the maximum likelihood (ML) tree (-In 
likelihood 544.123; 4909 trees examined) are generally congruent and differ from each 
other only in the position of the Cretean clade (Figs 2-3). This clade clusters separately 
in the NJ tree (Fig. 2) and groups with the sample of Selale (south Anatolia) in the ML 
tree (Fig. 3). However, both approaches revealed four main clades within Mesobuthus 
(Figs 2-3). The first clade is composed of the samples from the mainland of Greece 
(Vigla and Mathia) and island Rhodes (Petaloudes and Kolympia) and is found at a 
relatively low distance value. The two samples from Crete are found in a second 
independent clade, and the populations from Anatolia as a third clade. However, the 
Cyprus clade remains clearly separated from all other clades by a rather high tree length 
in both trees, and is confirmed as an offshot from the mainland. The bootstrap analysis 
supports four nodes of the trees with very high values (>90%), these are observed for 
the clades grouping the Cyprus samples (Tepebasi and Kantara), the central Anatolian 
samples (Avanos and Hacibectas), the Crete samples (Zakros and Vai), and the 
Androctonus species, respectively. The node between the Peloponnesian clade and the 
Cyprus clade is well supported (60% and 77%, respectively). Androctonus is confirmed 
as outgroup by a bootstrap value of 99%. However, the other nodes are weakly 
supported. The weakest support is found for the clade that contains the Cretean and 
Anatolian samples. Noteworthy, the outgroup species A. mauretanicus and A. crassi- 
cauda are separated by about the same genetic distance that is found among the samples 


A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 219 


of the Peloponnesus / Rhodes clade and the Anatolian clade, although the outgroup 
samples are treated as different species and are geographically separated by a large 
distance (approximatively 3000 km). 


Avanos 


Mesobuthus Anatolia 


Hacibectas 
Selale 
Zakros 

Vai Crete 
Vigla 


Mathia 


Peloponnesus 


Cyprus | 
Kantara 
96! Tepebasi 
Petaloudes 


Kolympia Rhodes 


Androctonus Urfa —— 


0.1 0 


91 Aourir Nei's D 


FIG. 2 
NJ tree based on Nei’s distance relating island and mainland populations of Mesobuthus inferred 


from 16 allozyme loci. A. crassicauda from Urfa (TR) and A. mauretanicus from Aourir (MA) 
were used as outgroups. 


TAXONOMY 


Mesobuthus cyprius Gantenbein & Kropf, sp. n. Figs 4-28; Tab. | 


Type material: Holotype: 1 male, Tepebasi, Northern Cyprus, 26. V. 1998, 
Natural History Museum Berne, Switzerland (NMBE). Paratypes (collected at the type locality): 
I male, 20. V. 1998, 1 female, 26. V. 1998, NMBE. 1 male, 20. V. 1998, 1 female, 26. V. 1998, 
Natural History Museum Basel, Switzerland; all specimens leg. A. Scholl (Berne, CH). 

Other material examined: Morphology: Mesobuthus cyprius sp. n.: 6 males, 
11 females from Tepebasi, I male, 1 female from Kantara, Cyprus. For a comparison, also 
specimens of Mesobuthus gibbosus from Rhodes, GR (31 specimens), Peloponnesus, GR (7), 
Crete, GR (20), Euboea, GR (1), Selale, TR (12) and central Anatolia, TR (3) were investigated. 

Allozyme study: Mesobuthus cyprius sp. n., Mesobuthus gibbosus, 
Androctonus mauretanicus, Androctonus crassicauda: See Table 1. 

Diagnosis: Mesobuthus cyprius Sp. n. can be distinguished unambiguously 
only by the shape of the basal lobes of the hemispermatophores. These are slender and 
acutely pointed teeth in Mesobuthus cyprius sp. n. (Figs 8-9), while in all populations of 
M. gibbosus examined they form scales with more or less distinct blunt tips (Figs 29- 


220 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


Vai 
99! Zakros Crete 
Selale 
Mesobuthus Avanos Anatolia 
Hacibectas 
4 
> Vigla 
Peloponnesos 
Mathia 


Kantara 
99 | Tepebasi Cyprus 


Petaloudes 
Kolympia Rhodes 
Androctonus Urfa 
89 Aourir 

freni 
0.1 0 

REML distance 

Fic. 3 


Rooted maximum likelihood (ML) tree of island and mainland populations of Mesobuthus. 


33: Vachon, 1948: fig. 1). Moreover, the hemispermatophore of the new species is 
considerably smaller than in M. gibbosus. 

Description: Measurements (in mm): No apparent size dimorphism, 
although females longer, and males with a longer metasoma. Total length (measured 
dorsally from anterior margin of carapace to tip of stinger, with telson in horizontal 
position): Males 45-55, females 45-60. Carapace length (measured dorsally along 
midline): males 5.0-5.7, females 5.0-6.0. Carapace width (maximum distance between 
posterio-lateral edges): males 5.5-6.0, females 5.5-6.6. Metasoma length: males 28-37, 
females 27-35 (measured dorsally from anterior margin of first segment to tip of 
stinger). 

Carapace (Figs 4, 5): trapezoid-shaped, colour pattern more or less distinct, 
especially lateral pigmented areas hardly visible in some specimens. Granulation: ante- 
rior median, central median and central lateral carinae distinct, granulae in remaining 
areas mostly weak. Anterior margin with transverse row of bristles. 

Mesosoma (Figs 5, 6): Tergites I-VI with a pattern of pigmented longitudinal 
stripes, i.e. one median stripe and two pairs of lateral stripes (one paramedian, one ectal 
pair). These stripes partly broken, especially on tergite 6, colour varying from dark grey 
to medium brown (only alcohol preserved material examined, colour pattern more 
distinct in juveniles than in adults, in some adults hardly visible). Interspace between 
median and paramedian stripe less than 1-1,5 times as wide as paramedian stripe at 
posterior margin of tergites IV-VI. Tergite VII with more or less pronounced median 


A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 22] 


3 
+ 59) 
7 


Fic. 4-6 


Mesobuthus cyprius sp. n. — 4. Female carapace, dorsal view. — 5. Male carapace and mesosoma, 
dorsal view. — 6. Sternum and genital opercula, ventral view. 


B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


(O) 
[59] 
(N°) 


keel bearing granula. Tergites only sporadically with bristles or without. Sternites 
smooth, at least some of them with white guanin spots: these spots sometimes distri- 
buted all over the ventral and occasionally also the dorsal surface. Sternum subtrian- 
gular, with deep median depression flanked by a pair of bristles (Fig. 6). Genital 
opercula with a few scattered bristles (Fig. 6). Pectines with 20-23 (females) or 26-30 
(males) teeth. 

Hemispermatophore (Figs 7-9): Long, slender, of the flagelliform type. Four 
lobes at truncal flexure: internal lobe biggest, with a small distal hook, close to the 
external lobe, partly embracing it. Basal lobe developed as a slender and acutely 
pointed tooth. 

Metasoma (Figs 10-12): Segments I-IV with 10 longitudinal keels formed by 
rows of tubercles; one additional pair of keels mediodorsally being indistinct and hardly 
visible (not illustrated). Fourth segment 1.79 — 2.44 times longer than high in males, 
1.54 — 1.58 times in females (length measured dorsally, height measured as distance 
between dorsal and ventral keel). Segment V granulate ventrally, with 3-4 blunt teeth 
laterally at distal edge. Telson with dark tip of stinger, granulate on ventral side, with 
several short bristles, one ventral pair of them rising from subaculeate tubercle. 

Chelicera (Figs 13-14): Tibia (nomenclature following Hjelle 1990) with two 
large bristles dorsaliy near distal-prolateral edge, three shorter translucent hairs (several 
more of them on movable finger) and a brush of fine hairs prolaterally. Fixed finger 
with two ventral and four dorsal teeth; medial and basal dorsal tooth fused to form a 
bicusp. Movable finger with five dorsal teeth, the two basal ones being the smallest; 
ventrally with three teeth, i.e. one distal and two medial ones. 

Pedipalp (Figs 15-23): Trichobothrial pattern constant, corresponding to type A 
of Vachon (1974). Femur a little shorter than patella, with 11 trichobothria, d2 
(nomenclature of trichobothria following Sissom, 1990) short and translucent (Fig. 17). 
Patella with two distinct tooth-like tubercles ventro-proximally and with 13 tricho- 
bothria, d2 short and translucent (Figs 18-19). Chela slender, males with a weak lobe on 
movable finger (not illustrated). 15 trichobothria present; esb, Esb and Eb3 short and 
translucent (Figs 20-21). Teeth on inner side of fingers arranged in a characteristic 
pattern, interspersed with short, stiff bristles, i.e. groups of two retrolateral and one 
prolateral bristle and several of them at the tip (Figs 22-23). 

Legs (Figs 24-28): increasing in length from leg | - 4. All legs with a pair of 
ventral pedal spurs distally on basitarsus. Proventral spur carrying a hair and a 
translucent tooth originating proximally on it (Fig. 28). Retroventral spur with a small 
tooth close to spur (Fig. 27). Legs 3 and 4 with a tibial spur distally (Fig. 26). Dorsal 
distitarsus with a conspicuous distal hair on an elevated base (Fig. 28). Apotele with 
three claws (Fig. 27). 

Remarks: Mesobuthus cyprius sp. n. differs from most populations of M. 
gibbosus by its pigmentation pattern on the mesosomal tergites, particularly by the 
width of the pale interspace between the median and the paramedian dark stripe which 
is less than 1-1,5 times the width of the latter at the posterior margin of tergites IV-VI. 
In most populations of M. gibbosus, the width of this interspace is at least more than 2 
times the width of the paramedian stripe which may even be absent. However, this 


i) 
(NO) 
(SS) 


A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 


0,5 mm 
> GE 
Se 


Se > A 


Fics 7-12 
Mesobuthus cyprius sp. n. — 7. Left hemispermatophore, total view; surrounding tissue of paraxial 
organ removed, except in distal part of flagellum. — 8, 9. Lobes at truncal flexure. — 10. 


Metasoma segments 4, 5, and telson of male, dorsal view. — 11. Ditto, lateral view. — 12. Ditto, 
ventral view. 


224 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


WA 
ZI I (1 A 
i LA 


Fic. 13-16 


Mesobuthus cyprius sp. n., male — 13. Right chelicera, ventral view; prolateral hair brush 
simplified. — 14. Ditto, dorsal view. — 15. Right pedipalp, ventral view. — 16. Ditto, dorsal view. 


A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 225 


DN 
ù SN EN alte ele 

Le.) a, NEE BZ 
oo, 20800 5608020 es 


Cesa = 
z Ge — Cai 0,5 mm 


22 


Fics 17-23 


Mesobuthus cyprius sp. n., male — 17. Right palpal femur, trichobothrial pattern, dorsal view. — 
18. Right palpal patella, trichobothrial pattern, dorsal view. — 19. Ditto, retrolateral view; arrow 
and asterisk in Figs 18, 19 indicate the same trichobothria. — 20. Right chela, trichobothrial 
pattern, proventral view. — 21. Ditto, retrodorsal view. — 22. Distal part of fixed finger of right 
chela, inner side. — 23. Tip of movable finger of right chela, inner side. 


226 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


Fics 24-28 


Mesobuthus cyprius sp. n., male legs. — 24. Left leg I, prolateral view. — 25. Ditto, retrolateral 
view. — 26. Left leg IV, prolateral view. — 27. Left tarsus I, ventral view; hairs omitted. — 28. 
Ditto, dorsal view. 


A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS DDT 


2 
2. 


Bar 30. Le 


0,5 mm 


IEZZO 


BO as 33 


Fics 29-33 


Mesobuthus gibbosus, lobes at truncal flexure of left hemispermatophore in specimens from 
different regions; same magnification as in Figs 8-9. — 29. Euboea (GR). — 30. Rhodes, Kolympia 
(GR). — 31. Rhodes, Monolithos (GR). — 32, 33. Selale (TR). 


228 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


distinction is not reliable. On the one hand the mesosomal pattern is hardly visible in 
some animals, on the other hand specimens of M. gibbosus from the Peloponnesian 
peninsula (Mathia) and from Crete can also show this pigmentation pattern. The 
Peloponnesian individuals of M. gibbosus differ by a characteristic row of bristles on 
the mesosomal tergites, which cannot be found in Mesobuthus cyprius sp. n. There, 
only some scattered bristles, if any at all, do occur. Some of the Cretean specimens of 
M. gibbosus examined cannot be distinguished from M. cyprius sp. n. by means of 
external morphology, because they possess a similar pigmentation pattern and lack 
mesosomal bristle rows. 


DISCUSSION 


The allozyme data generally revealed a low genetic variability (Table 1), as 
already observed in earlier studies on scorpions of the family Euscorpiidae (Stockwell 
1992; Gantenbein ef al., 1998, 1999). One exception is the population of M. gibbosus 
from the island of Rhodes which shows a higher level of heterozygosity as compared to 
the other Mesobuthus populations. This is mainly due to three gene loci (Ark, Gtdh, 
Sod) (Table 1). Specimens of M. gibbosus from Rhodes that are heterozygous at these 
three loci possess one allele, which is elsewhere only found in the samples from the 
Peloponnesus, and another allele, which is common in the Cretean and Anatolian 
populations. We speculate that this pattern indicates hybridization between western 
Aegean populations and eastern Aegean populations on the island of Rhodes. This 
hybridization may have resulted from secondary contact between the authochtonous 
Rhodes population and an introduced population of Greek origin. 

The allozyme gene frequencies indicate that there are different population 
groups within Mesobuthus. In the NJ and ML tree, the Cyprus clade is separated from 
the other clades by a long branch (Figs 2-3). Both approaches clearly underline that the 
Cyprus scorpions have diverged considerably. Since the Biological Species Concept 
(BSC) (Mayr 1942) can hardly be applied to allopatric populations, it remains unsolved 
if the Cyprus population represents a separate species in this sense. Consequently, there 
are no objective arguments for the taxonomic rank of this population. However, our 
expectation that this island population remained isolated from the mainland populations 
since the colonisation 5.2 Myrs ago is supported by a high genetic divergence of the 
Cyprus clade and by the noninterchangeable morphology of the hemispermatophores. 
Taking these two arguments into account we, therefore, designate to the Cyprus 
population the rank of a species. 

It is remarkable that M. cyprius sp. n. appears genetically distinct and highly 
differentiated, but can hardly be characterised by means of classical morphology. Such 
‘hidden’ taxa are well known from other animal groups (e. g. mosquitos, polychaetes, 
myriapodes) (Narang ef al., 1989), especially on islands. The only characters that can 
be used for a morphological distinction are found in the hemispermatophores, in the 
mesosomal pigmentation pattern and in a lack of mesosomal bristle rows. The last 
character is a negative trait and presumably represents a plesiomorphic state as 
compared to most populations of M. gibbosus. The bristle row was found in specimens 


A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 229 


of M. gibbosus from the Peloponnesus (Mathia) and from Rhodes (Petaloudes). The 
pigmentation pattern seems to be quite useful, because we found only limited variation 
there (but compare paragraph ‘remarks’ above). The taxonomic value of the pigmen- 
tation pattern was underlined by Lourenco (1983), who claimed that species with poly- 
morphic colouration or pigmentation are an exception among buthids. As a conclusion, 
only a single reliable distinctive morphological character of M. cyprius sp. n. remains 
discriminant, 1.e. the shape of the basal lobes of the hemispermatophores. 

Based on a single male, Schenkel (1947) described a new variety of M. gibbosus 
from central Anatolia (collected along the road between Sivas and Amasya, Turkey). 
Kinzelbach (1975) elevated Schenkel’s variety to subspecies rank. He distinguished 
M. g. gibbosus (Brullé, 1832) in the south of the Balkan Peninsula, Northern Sporades, 
Cyclades, from M. g. anatolicus (Schenkel 1947) living in Crete, Anatolia, and Cyprus, 
using differences in the number of pectinal teeth as the main character. Furthermore, he 
argued that these two subspecies were separated since the Miocene (12-25 Myrs BP). 
Kritscher (1993) refuted Kinzelbach’s arguments for a separate subspecies anatolicus 
and rejected this taxon. The allozyme data now partially support the existence of two 
subspecies in Kinzelbach’s sense. The Greek mainland populations are genetically 
slightly distinct from the Crete-Anatolian population group (Figs 2, 3). However, the 
Cyprus population (= Mesobuthus cyprius sp. n.) is genetically considerably more 
differentiated from all others, as discussed above. This fact contradicts the existence of 
M. g. anatolicus on Cyprus as proposed by Kinzelbach. 


ACKNOWLEDGEMENTS 


Rica and Andreas Quensel, Ahmet Ylzaz and Keco Kutlay, Girne (Northern 
Cyprus) provided logistics for field work and collecting in Northern Cyprus and 
Turkey. Iasmi Stathi, Matt Braunwalder and Peter Schwendinger (Muséum d’histoire 
naturelle, Geneve, CH) contributed specimens from Crete. Field trips were financially 
supported by the Dr. Karl Bretscher-Foundation Berne. Beatrice Liischer and Lilian 
Beer assisted in the allozyme analysis. The comments of Victor Fet and two anonymous 
reviewers greatly improved the final version of the manuscript. 


REFERENCES 


AYALA, F. J., POWELL, J. R., TRACEY, M. L., Mourao, C. A. & PEREZ-SALAS, S. 1972. Enzyme 
variability in the Drosophila willistoni group. IV. Genic variation in natural populations 
of Drosophila willistoni. Genetics 70: 113-139. 

BALDWIN, B. G. & SANDERSON, M. J. 1998. Age and rate of diversification of the Hawaiian 
silversword alliance (Compositae). Science 95: 9402-9406. 

BEERLI, P., Hotz, H. & U7ZELL, H. 1996. Geologically dated sea barriers calibrate a protein clock 
for Aegean Water frogs. Evolution 50: 1676-1687. 

BENTON, T. G. 1991. The life history of Euscorpius flavicaudis (Scorpiones, Chactidae). Journal 
of Arachnology 19: 105-110. 

Carson, H. L. & TEMPLETON, A. R. 1984. Genetic revolutions in relation to speciation pheno- 


mena: The founding of new populations. Annual Review of Ecology and Systematics 15: 
97-131. 


230 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


CAVALLI-SFORZA, L. L. & EDWARDS, A. W. F. 1967. Phylogenetic analysis: Models and esti- 
mation procedures. Evolution 32: 550-570. 

CLAYTON, J. W. & TRETIAK, D. N. 1972. Amine-citrate buffers for pH control in starch gel 
electrophoresis. Journal of Fisheries Research Board of Canada 29: 1169-1172. 

CRUCITTI, P. 1993. Some topics on distribution patterns of the genus Mesobuthus in the Near East 
based on ecological data (Scorpiones: Buthidae). Biologia Gallo-helenica 20(1): 69-74. 
(5° Congrès international sur la zoogéographie et l'écologie de la Grèce et des régions 
avoisinantes - Iraklion, Avril 1990). 

ESTOUP, A., SOLIGNAC, M., CORNUET, J.-M., GOUDET, J. & SCHOLL, A. 1996. Genetic differentia- 
tion of continental and island populations of Bombus terrestris (Hymenoptera: Apidae) in 
Europe. Molecular Ecology 5: 19-31. 

FELSENSTEIN, J. 1973. Maximum-likelihood estimation of evolutionary trees from continuous 
characters. American Journal of Human Genetics 25: 471-492. 

FELSENSTEIN, J. 1981. Evolutionary trees from gene frequencies and quantitative characters: 
Finding maximum likelihood estimates. Evolution 35: 1229-1242. 

FELSENSTEIN, J. 1984. Distance methods for inferring phylogenies: a justification. Evolution 38: 
16-24. 

FELSENSTEIN, J. 1995. PHYLIP (Phylogeny Inference Package), Version 3.57c. Seattle, University 
of Washington. 

FET, V. 1986. Notes on some Euscorpius (Scorpiones, Chactidae) from Greece and Turkey. 
Rivista del Museo civico di scienze naturali "Enrico Caffi" (Bergamo) 9: 3-11. 

FET, V. & RECHKIN, D. V. 1989. Scorpion trichobothriotaxy: a principal component analysis. 
Rivista del Museo civico di scienze naturali "Enrico Caffi" (Bergamo) 14: 191-206. 

GANTENBEIN, B., BUCHI, L., BRAUNWALDER, M. E. & SCHOLL, A. 1998. The genetic population 
structure of Euscorpius germanus (C. L. Koch) (Scorpiones: Chactidae) in Switzerland 
(pp. 33-40). In: Selden, P. A. (ed). Proceedings of the 17th European Colloquium of 
Arachnology. Edinburgh 1997, 14-18 July, 1997. 

GANTENBEIN, B., FET, V., LARGIADER, C. R. & SCHOLL, A. 1999. First DNA phylogeny of 
Euscorpius Thorell, 1876 (Scorpiones, Euscorpiidae) and its bearing on taxonomy and 
biogeography of this genus. Biogeographica 75: 49-65. 

GILLESPIE, R. G., RIVERA, M. A. & GARB, J. E. 1998. Sun, surf and spiders: taxonomy and phylo- 
geography of Hawaiian Araneae (pp. 41-51. /n: Selden, P. A. (ed). Proceedings of the 
17th European Colloquium of Arachnology. Edinburgh 1997, 14-18 July, 1997. 

GOYFFON, M. 1992. Le rôle de l’homme dans l’expansion territorial de quelques espèces de 
scorpions. Bulletin de la Société zoologique de France 177: 15-19. 

GRANT, P. R. 1998. Speciation (pp. 83-101). In: Grant, P. R. (ed.). Evolution on islands. Oxford 
University Press, Oxford, 319 pp. 

GRUBER, J. 1963. Ergebnisse der von Dr. O. Paget und Dr. E. Kritscher auf Rhodos durch- 
geführten zoologischen Exkursionen. VII. Scorpiones und Opiliones. Annalen des Natur- 
historischen Museums in Wien 66: 307-316. 

GRUBER, J. 1966. Ergebnisse der von Dr. O. Paget und Dr. E. Kritscher auf Rhodos durch- 
geführten zoologischen Exkursionen. VII. Scorpiones und Opiliones (2.Teil). Annalen 
des Naturhistorischen Museums in Wien 69: 423-426. 

HOLLOCHER, H. 1998. Island hopping in Drosophila: genetic patterns and speciation mechanisms 
(pp. 124-141). Zn: Evolution on islands. Grant, P. R. (ed.). Oxford University Press, 
Oxford, 319 pp. 

HIELLE, J. T. 1990. Anatomy and Morphology. Chapter 2 (pp. 9-63). In: Biology of Scorpions. 
Polis, G. A. (ed.). Stanford University Press, Stanford, California, 587 pp. 

Hsü, K. J. 1972. Origin of the saline giants: a critical review after the discovery of the Medi- 
terranean Evaporite. Earth-Science Reviews 8: 371-396. 


A NEW BUTHID TAXON ON THE ISLAND OF CYPRUS 23] 


Hsu, K. J., MONTADERT, L., BERNOULLI, D., CITA, M. B., ERICKSON, A., GARRISON, R. E., KIDD, 
R. B., MELIERES, F., MÜLLER, C. & WRIGHT, R. 1977. History of the Mediterranean 
salinity crisis. Nature 267: 399-403. 

KINZELBACH, R. 1975. Die Skorpione der Ägäis. Beiträge zur Systematik, Phylogenie und Bio- 
geographie. Zoologische Jahrbücher, Abtheilung für Systematik 102(1): 12-50. 

KINZELBACH, R. 1982. Die Skorpionssammlung des Naturhistorischen Museums der Stadt Mainz. 
- Teil I. Europa und Anatolien. Mainzer naturwissenschaftiches Archiv 20: 49-66. 

KINZELBACH, R. 1984. Die Skorpionssammlung des Naturhistorischen Museums der Stadt Mainz. 
- Teil II: Vorderasien. Mainzer naturwissenschaftiches Archiv 22: 97-106. 

KINZELBACH, R. 1985. Vorderer Orient. Skorpione (Arachnida: Scorpiones). Tübinger Atlas des 
Vorderen Orients (TAVO) Karte Nr. A VI 14.2. 

KRITSCHER, E. 1993. Ein Beitrag zur Verbreitung der Skorpione im östlichen Mittelmeerraum. 
Annalen des Naturhistorischen Museums in Wien (B) 94/95: 377-391. 

LOURENCO, W. R. 1983. Importance de la pigmentation dans l’étude taxonomique des Buthidae 
néotropicaux (Arachnida, Scorpiones). Bulletin du Museum national d’histoire naturelle 
(Paris) 5: 611-618. 

Mayr, E. 1942. Systematics and the origin of species. Columbia University Press, New York, 

334 pp. 

MICHALIS, K. & KATTOULAS, M. 1981. A systematic, ecological, zoogeographical and biometrical 

study of the scorpions of the Peloponnesus. Opuscula Zoologica (Budapest) X VII-X VIII: 

107-112: 

MICHALIS, K. & DOLKERAS, P. 1989. Beitrag zur Kenntnis der Skorpione Thessaliens und Epirus 
(Nordgriechenland). Entomologische Mitteilungen aus dem Zoologischen Museum Ham- 
burg 9: 259-270. 

MURPHY, R. W., SITES, J. W., BUTH, D. G. & HAUFLER, C. H. 1996. Proteins: Isozyme electro- 
phoresis. Chapter 4, pp. 51-120. /n: Hillis, D. M., Moritz, C. & Mable, B. K. (eds). Mole- 
cular systematics. 2™ edition. Sinauer Assoc. Inc., Massachusetts, 655 pp. 

NARANG, S. K., KAISER, P. E. & SEAWRIGHT, J. A. 1989. Dichotomous electrophoretic key for 

idenification of sibling species A, B and C of the Anopheles quadrimaculatus complex 

(Diptera: Culicidae). Journal of Medical Entomology 26: 94-99. 

NEI, M. 1972. Genetic distance between populations. American Naturalist 106: 283-292. 

NEI, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of 

individuals. Genetics 83: 583-590. 

NEI, M. 1987. Genetic distance and molecular phylogeny. Chapter 8 (pp. 193-215). Jn: Nils, N. & 
Utter, F. (eds). Population genetics & fishery. University of Washington Press, Seattle, 
417 pp. 

PAGE, R. D. M. & HOLMES, E. C. 1998. Molecular evolution. A phylogenetic approach. Chapter 5 
(pp. 172-227). Blackwell Science, London, 347 pp. 

Saitou, N. & NEI., M. 1987. The neighbor-joining method: a new method for reconstructing 
phylogenetic trees. Molecular Biology and Evolution 4: 406-425. 


SCHENKEL, E. 1947. Einige Mitteilungen über Spinnentiere. Revue suisse de Zoologie 54: 1-16. 

SissoM, W. D. 1990. Systematics, biogeography and paleontology. Chapter 3 (pp. 64-160). In: 
Polis G. A., (ed.). Biology of Scorpions. Stanford University Press, Stanford, California, 
587 pp. 

STOCKWELL, S. A. 1992. Systematic observations on North American Scorpionida with a key and 
checklist of the families and genera. Journal of Medical Entomology 29: 407-422. 
TEMPLETON, A. R. 1980. The theory of speciation via the founder principle. Genetics 91: 1011- 

1038. 
TEMPLETON, A. R. 1981. Mechanisms of speciation - a population genetic approach. Annual 
Review of Ecology and Systematics 12: 23-48. 


232 B. GANTENBEIN, C. KROPF, C. R. LARGIADER & A. SCHOLL 


TOLUNAY, M. A. 1959. Zur Verbreitung der Skorpione in der Türkei. Zeitschrift für angewandte 
Entomologie 43: 366-370. 


TOSCANO-GADEA, C. A. 1998. Euscorpius flavicaudis (de Geer, 1778) in Uraguay: First record 
from the New World. Newsletter of the British arachnological Society 81: 6. 


VACHON, M. 1947a. Remarques préliminaires sur le faune des Scorpions de Turquie. Bulletin du 
Museum national d’histoire naturelle (Paris) 19: 161-164. 


VACHON, M. 1947b. Répartition et origine des scorpions de Turquie. Comptes rendus des séances 

de la Société de biogéographie 206: 26-27. 

VACHON, M. 1948. Scorpions récoltés dans l'île de Crête par Mr. le Docteur Otto von Wettstein. 
Annalen des Naturhistorischen Museums in Wien 56: 60-69. 


VACHON, M. 1966. Liste des Scorpions connus en Égypte, Arabie, Israël, Liban, Syrie, Jordanie, 
Turquie, Irak, Iran. Toxicon 4: 209-218. 

VACHON, M. 1974. Etude des caractères utilisés pour classer les familles et les genres de 
Scorpions (Arachnides). 1. La trichobothriotaxie en Arachnologie, sigles trichobothriaux 
et types de trichobothriotaxie chez les Scorpions. Bulletin du Museum national d'histoire 
naturelle (Paris), 140 (104): 857-958. 


VACHON, M. 1975. Sur l'utilisation de la trichobothriotaxie du bras des pédipalpes des Scorpions 
(Arachnides) dans le classement des genres de famille des Buthidae Simon. Comptes 
rendus hebdomadaires des séances de l'Académie des sciences (D) 281: 1597-1599. 


VACHON, M. 1976. Formules permettant de localiser l'asymétrie numérique de certains caractères 
trichobothriotaxiques utilisés dans la classification spécifique ou intraspécifique chez les 
Scorpions (Arachnides). Comptes rendus hebdomadaires des séances de l'Académie des 
sciences (D), 282: 1681-1684. 

VACHON, M. & KINZELBACH, R. 1987. On the taxonomy and distribution of the scorpions of the 
Middle East (pp. 91-103). /n: Krupp, F., Schneider, W. & Kinzelbach, R. (eds). Pro- 
ceedings of the Symposium on the Fauna and Zoogeography of the Middle East, Mainz, 
1985. Beihefte zum Tübinger Atlas des vorderen Orients, Reihe A (28) (Naturwissen- 
schaften). 

VACHON, M. & ABE, T. 1988. Colonization of the Krakatau Islands (Indonesia) by scorpions. 
Acta Arachnologica 37: 23-32. 

WERNER, F. 1928. Beiträge zur Kenntnis der Fauna Griechenlands, namentlich der ägäischen 
Inseln. Sitzungsberichte der Akademie der Wissenchaften in Wien, 137: 283-295. 
WERNER, F. 1937. Beiträge zur Kenntnis der Tierwelt des Peloponnes der Inseln Kythira und 
Euboea sowie der kleinen Inseln im Saronischen Golf. Sirzungsberichte der Akademie der 

Wissenschaften in Wien 146: 135-153. i 

WERNER, F. 1938. Ergebnisse der achten zoologischen Forschungsreise nach Griechenland 
(Euboea, Tinos, Skiathos, Thasos usw.). Sitzungsberichte der Akademie der Wissen- 
schaften in Wien 147: 151-173. 

WIDMER, A., SCHMID-HEMPEL, P., Estoup, A. & SCHOLL, A. 1998. Population genetic structure 
and colonization history of Bombus terrestris s.I. (Hymenoptera: Apidae) from the 
Canary Islands and Madeira. Heredity 81: 563-572. 


REVUE SUISSE DE ZOOLOGIE 


Tome 107 — Fascicule | 


JUVARA-BALS, Ilinca & Vojciech WITALINSKI. Description of five new 
species of Holoparasitus s. str. with redescription of H. apenninorum 
(Berlese, 1906) and H. cultriger (Berlese, 1906) from Italy and Spain 
(AcangGamasida Para side NE PER RO 


SiLva, Carla Maria Menegola da & Beatriz MOTHES. Three new species of 
Geodia Lamarck, 1815 (Porifera, Demospongiae) from the bathyal 
depths off Brazilian coast, Southwestern Atlantic................. 


MAHUNKA, Sandor & Luise MAHUNKA-Papp. Oribatids from Switzerland III 
(Acari: Oribatida: Oppiidae | and Quadroppiidae). (Acarologica Ge- 
GV CHEST ODN CNN a I Us 
GIL DE PERTIERRA, Alicia A. & Alain de CHAMBRIER. Rudolphiella szidati 
sp. n. (Proteocephalidea: Monticelliidae, Rudolphiellinae) parasite of 
Luciopimelodus pati (Valenciennes, 1840) (Pisces: Pimelodidae) from 
Argentina with new observations on Rudolphiella lobosa (Riggenbach, 
1006): a N en CE E ce Gene 


BENJAMIN, Suresh P. & Rudy JOCQUÉ. Two new species of the genus 
Sujjasia trom Sr Lanka (Araneae: Zodariidae). . dr Re 


LANG, Claude. Diversité du zoobenthos dans 47 rivières du canton de Vaud: 
temaance yO SIERO Re N RE ES 


VIT, Stanislav. Contribution a la connaissance de la famille Eucinetidae 
(COEO OS) ARS EE eno DR PERU ev Wee Cia 


BÄNZIGER, Ruth. Spatio-temporal distribution of size classes and larval 
instars of aquatic insects (Ephemeroptera, Trichoptera and Leopi- 
doptera) in a Potamogeton pectinatus L. bed (Lake Geneva, Switzer- 
Fann) reenact ee Wee el NG NER SINE NT Vogt RELA, 


BESUCHET, Claude & Stanislav VIT. Les Nanophthalmus Motschulsky 
eEurope (Coleoptera Scydmacnidac))a ri 220, nen, 


HEYD, Andreas & Wolfgang PFEIFFER. Uber die Lauterzeugung der Welse 
(Siluroidei, Ostariophysi, Teleostei) und ihren Zusammenhang mit der 
Biyloseniesund der Schreckrealsti ona 2... nun. on: 


GANTENBEIN, Banjamin, Christian KRoPF, Carlo Rodolfo LARGIADER & 
Adolf SCHOLL. Molecular and morphological evidence for the presence 
of a new Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus 


Pages 


81-95 


97-106 


107-122 


123-138 


139-151 


153-163 


165-211 


213232 


REVUE SUISSE DE ZOOLOGIE 


Volume 107 — Number 1 


JUVARA-BALS, Ilinca & Vojciech WITALINSKI. Description of five new 
species of Holoparasitus s. str. with redescription of H. apenninorum 
(Berlese, 1906) and H. cultriger (Berlese, 1906) from Italy and Spain 
(Sean Gamasıda-Barasıtidae).r or RR 


SILVA, Carla Maria Menegola da & Beatriz MOTHES. Three new species of 
Geodia Lamarck, 1815 (Porifera, Demospongiae) from the bathyal 
depths off Brazilian coast, Southwestern Atlantic................. 


MAHUNKA, Sandor & Luise MAHUNKA-Papp. Oribatids from Switzerland III 
(Acari: Oribatida: Oppiidae | and Quadroppiidae). (Acarologica Ge- 
Haven ar NEIL) wipes ewe oe nce: Ne alto cok ed er 


GIL DE PERTIERRA, Alicia A. & Alain de CHAMBRIER. Rudolphiella szidati 
sp. n. (Proteocephalidea: Monticelliidae, Rudolphiellinae) parasite of 
Luciopimelodus pati (Valenciennes, 1840) (Pisces: Pimelodidae) from 
Argentina with new observations on Rudolphiella lobosa (Riggenbach, 
DS OS) EMEA O: IR CARTIER, et RUES RC NES, (REA eee 


BENJAMIN, Suresh P. & Rudy JOCQUÉ. Two new species of the genus 
Suffasia from Sri Lanka (Araneae: Zodariidae). . ................ 


LANG, Claude. Diversity of zoobenthos in 47 rivers of western Switzerland: 
HEIISIENIITÄTLENIR RM IONI E 


Vit, Stanislav. Contribution to the knowledge of Eucinetidae (Coleoptera). . 


BANZIGER, Ruth. Spatio-temporal distribution of size classes and larval 
instars of aquatic insects (Ephemeroptera, Trichoptera and Leopi- 
doptera) in a Potamogeton pectinatus L. bed (Lake Geneva, Switzer- 
lands ra LR US kien a er E AUTOMATE. De RARE 

BESUCHET, Claude & Stanislav VIT. European species of the genus Nanop- 
thalmus Motschulsky (Coleoptera, Scydmaenidae)............... 

HEYD, Andreas & Wolfgang PFEIFFER. Sound production in catfish (Siluro- 
idei, Ostariophysi, Teleostei) and its relationship to phylogeny and 
fright means BOC MGA DIR ON SERIE EB SER 

GANTENBEIN, Banjamin, Christian KROPF, Carlo Rodolfo LARGIADER & 
Adolf SCHOLL. Molecular and morphological evidence for the presence 
of anew Buthid taxon (Scorpiones: Buthidae) on the Island of Cyprus 


Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX 


Pages 


3-30 


31-48 


49-79 


81-95 


97-106 


107-122 
123-138 


139-151 


153-163 


165-211 


213-232 


PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE 


CATALOGUE DES INVERTEBRES DE LA SUISSE, N9S 1-17 (1908-1926)... serie Fr. 285.— 


(prix des fascicules sur demande) 
REMIUEIDEIPÄNEOBIOLOGIEN FT. nno Echange ou par fascicule Fr. 
LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris) ....... par fascicule Fr. 


COLLEMBOLENFAUNA EUROPAS 

von H. Gisin, 312 Seiten, 554 Abbildungen, 1960 (Nachdruck, 1984) ............. Fr 
THE EUROPEAN PROTURA THEIR TAXONOMY, ECOLOGY AND 

DISTRIBUTION WITH KEYS FOR DETERMINATION 

DYAANOSER 34 pages llifioures intex MOTS ee ee. Fr. 
CLASSIFICATION OF THE DIPLOPODA 

par Richard L. HOFFMAN, 237 pages, 1979 
LES OISEAUX NICHEURS DU CANTON DE GENEVE 


par P. GEROUDET, C. GUEX et M. MAIRE 
SH lgpacess mombneusesicanlesjeti Sunessal O83. ely sia A SN I Fr. 


CATALOGUE COMMENTE DES TYPES D'ECHINODERMES ACTUELS 
CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES, 
SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ 
A LA CONNAISSANCE DES ECHINODERMES ACTUELS 
pa MichelyANGOUXHOV paces: Lilplanchess ISSN 2 u cae ces en Er. 


RADULAS DE GASTEROPODES LITTORAUX DE LA MANCHE 

(COTENTIN-BAIE DE SEINE, FRANCE) 

Paty EINE = WUESD ct ke MAREDA 62:pages, LINE Ten. Fr. 
GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN; 

SHELLS AND RADULAS 

DYSARINER es WiWESmianduke MAREDA OOD. ee Fr. 
O. SCHMIDT SPONGE CATALOGUE 

par R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 pages, 49 plates, 1992 .......... Fr. 
ATLAS DE REPARTITION DES AMPHIBIENS 

ET REPTILES DU CANTON DE GENEVE 

Paw Ae KEPLER, Vi AELLENIet V.. MAHNERT, 48ipages, 1993 FU re Fr. 
THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: 

A DOCUMENTED FAUNAL LIST 

par Yves Pine, SUDAN ISLE A sol du ae Er: 
NOTICE SUR LES COLLECTIONS MALACOLOGIQUES 

DU MUSEUM D’HISTOIRE NATURELLE DE GENEVE 

Pawlean-@laudei@RınnıEzsA9 pages MISERA Fr. 
PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS 

OF ARACHNOLOGY, Geneva 1995 (ed. Volker MAHNERT), 720 pages, 1996 ....... Fr. 


CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) 
(Instrumenta Biodiversitatis I) par Ivan LOBL, 192 pages, 1997 .................... Fr. 


CATALOGUE SYNONYMIQUE ET GÉOGRAPHIQUE DES SYRPHIDAE (DIPTERA) 

DE LA REGION AFROTROPICALE 

(Instrumenta Biodiversitatis II) par Henri G. DIRICKX, 188 pages, 1998 ............ Fr. 
A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE 

WEST PALAEARCTIC REGION 


(Instrumenta Biodiversitatis III) par Stanley BOWESTEAD, 204 pages, 1999 ............ Fr. 


THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE 
SOKOTRA ARCHIPELAGO 
(Instrumenta Biodiversitatis IV) par Beat SCHATTI & Alain DESVOIGNES, 
178 pages, 1999 


35.— 
35.— 


30. — 


30.— 


30.— 


30.— 


40.— 


22.— 


r. 70.— 


ins 


796 Ne 
VE LAURE 


> AGG af 


Volume 107 - Number 1 - 2000 
Revue suisse de Zoologie: Instructions to Authors 


The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific 
results based on the collections of the Muséum d’histoire naturelle, Geneva. Submission of a manuscript implies 
that it has been approved by all named authors, that it reports their unpublished work and that it is not being 
considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of 
colour plates and large manuscripts. All papers are refereed by experts. 

In order to facilitate publication and avoid delays authors should follow the /nstructions to Authors and refer to 
a current number of R.S.Z. for acceptable style and format. Manuscripts not conforming with these directives are 
liable to be returned to the authors. Papers may be written in French, German, Italian and English. Authors should 
aim to communicate ideas and information clearly and concisely. Authors not writing in their native language should 
pay particular attention to the linguistic quality of the text. 

Manuscripts must be typed, or printed (high quality printing, if possible by a laser-printer), on one side only 
and double-spaced, on A4 (210 x 297 mm) or equivalent paper and all pages should be numbered. All margins 
must be at least 25 mm wide. Authors must submit one original and two copies, including tables and figures, in 
final fully corrected form, and are expected to retain another copy. 

We encourage authors to submit the text on a diskette (3,5’’, Macintosh or IBM compatible, with “Microsoft 
Word” or similar programmes). The text should be in roman (standard) type face throughout, including headings, 
except genus and species names which should be formatted in italics (or underlined with pencil); bold, small 
capitals, large capitals and other type faces should not be used. Footnotes and cross-references by page should be 
avoided. 

Papers should conform to the following general layout: 

Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces, 
name(s) in full and surname(s) of author(s), and full address(es). 

Abstract. The abstract is in English, composed of the title and a short text of up to 200 words. It should 
summarise the contents and conclusions of the paper. The abstract is followed by less than 10 key-words, 
separated by hyphens, which are suitable for indexing. 

Introduction. A short introduction to the background and the reasons for the work. 

Materials and methods. Sufficient experimental details must be given to enable other workers to repeat the 
work. The full binominal name should be given for all organisms. The International Code of Zoological Nomen- 
clature must be strictly followed. Cite the authors of species on their first mention. 

Results. These should be concise and should not include methods or discussion. Text, tables and figures should 
not duplicate the same information. New taxa must be distinguished from related taxa. The abbreviations gen. n., sp. 
n., syn. n. and comb. n. should be used to distinguish all new taxa, synonymies or combinations. Primary types must 
be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed by 
synonyms, material examined and distribution, description and comments. All material examined should be listed in 
similar, compact and easily intelligible format; the information should be in the same language as the text. Sex 
symbols should be used rather than “male” and “female”. 

Discussion. This should not be excessive and should not repeat results nor contain new information, but 
should emphasize the significance and relevance of the results reported. 

References. The Harvard System must be used for the citation of references in the text, e.g. White & Green 
(1995) or (White & Green, 1995). For references with three and more authors the form Brown et al. should be 
used. Authors’ names should not be written in capitals. The list of references must include all publications cited in 
the text but only these. References must be listed in alphabetical order of authors, and both the title and name of 
the journal must be given in full in the following style (italics can be formatted by the author): 

Penard, E. 1888. Recherches sur le Ceratium macroceros. Thése, Genéve, 43 pp. 

Penard, E. 1889. Etudes sur quelques Héliozoaires d’eau douce. Archives de Biologie 9: 1-61. 

Mertens, R. & Wermuth, H. 1960. Die Amphibien und Reptilien Europas, Kramer, Frankfurt am Main, XI + 264 pp. 
Handley, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: Wenzel R. L. & Tipton, V. J. (eds). 

Ectoparasites of Panama. Field Museum of Natural History, Chicago, XII + 861 pp. 

References should not be interspaced and, in the case of several papers by the same author, the name has to be 
repeated for each reference. 

Tables. These should be self-explanatory, with the title at the top organised to fit 122 x 180 mm. Each table 
should by typed, double spaced, on a separate page and numbered consecutively and its position indicated in the text. 

Figures. These may be line drawings or half tones and all should be numbered consecutively, and their position 
indicated in the text. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and 
lettering should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors 
should refrain from mixing the drawings and half tones. Original drawings will not be returned automatically. The 
Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents. If electronically 
scanned figures are submitted on diskettes, this should be clearly indicated on the print-out enclosed with the 
manuscript. 

Legends to figures. These should be typed in numerical order on a separate sheet. 

Proofs. Page proofs only are supplied, and authors may be charged for alterations (other than printer’s errors) if 
they are numerous. 

‘Offprints. The authors receive totally 25 offprints free of charge; more copies may be ordered at current prices 
when proofs are returned. 

Correspondence. All correspondence should be addressed to 


Revue suisse de Zoologie 
Muséum d'histoire naturelle 
CP 6434 
CH-1211 Genève 6 
Switzerland. 
Phone: +41 22 418 63 33 - Fax +41 22 418 63 01 
e-mail: volker.mahnert @ mhn.ville-ge.ch 
Home page RSZ: http://www. ville-ge.ch/musinfo/mhng/page/rsz.htm 


N 


11901002Z 40 1SSINS INMAJA 


ADOTOOZ 40 IVNAYNOÏ SSIMS 


rd 


)00 ISSN 0035 - 418 X 


REVUE SUISSE DE ZOOLOGIE 


TOME 107 — FASCICULE 2 


Publication subventionnee par: 


ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN 
VILLE DE GENEVE 
SOCIETE SUISSE DE ZOOLOGIE 


VOLKER MAHNERT 
Directeur du Muséum d'histoire naturelle de Genève 


MANUEL RUEDI 
Chargé de recherche au Muséum d histoire naturelle de Genève 


CHARLES LIENHARD 
Chargé de recherche au Muséum d’ histoire naturelle de Genève 


Comité de lecture 


Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Geneve et de représentants des Instituts de zoologie des universités 
suisses. 

Les manuscrits sont soumis a des experts d’institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 
comparée, physiologie. 

Administration 


MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 


Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 


PRIX DE L'ABONNEMENT: 


SUISSE Fr. 225.— UNION POSTALE Fr. 230.— 


(en francs suisses) 


Les demandes d'abonnement doivent être adressées 
à la rédaction de la Revue suisse de Zoologie, 
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 


ANNALES 


de la 

SOCIETE SUISSE DE ZOOLOGIE 

et du 

MUSEUM D'HISTOIRE NATURELLE 
de la Ville de Genéve 


tome 107 
fascicule 2 
2000 


E < 
kl GENEVE JUIN 2000 ISSN 0035 - 418 X 


SWISS JOURNAL OF ZOOLOGY 


REVUE SUISSE DE ZOOLOGIE 


REVUE SUISSE DE ZOOLOGIE 


TOME 107 — FASCICULE 2 


Publication subventionnee par: 


ACADEMIE SUISSE DES SCIENCES NATURELLES ASSN 
VILLE DE GENEVE 
SOCIETE SUISSE DE ZOOLOGIE 


VOLKER MAHNERT 
Directeur du Muséum d'histoire naturelle de Genéve 


MANUEL RUEDI 
Chargé de recherche au Muséum d'histoire naturelle de Genève 


CHARLES LIENHARD 
Charge de recherche au Muséum d’histoire naturelle de Geneve 


Comité de lecture 


Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des Instituts de zoologie des universités 


suisses. 


Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 


sujet étudié. 


La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 


comparée, physiologie. 


Administration 


MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 


Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 


PRIX DE L'ABONNEMENT: 


SUISSE Fr. 225.— UNION POSTALE Fr 230 


(en francs suisses) 


| 
| Les demandes d'abonnement doivent étre adressées 
| à la rédaction de la Revue suisse de Zoologie, 
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 


REVUE SUISSE DE ZOOLOGIE 107 (2): 233-243; juin 2000 


Etat trophique du lac de Morat indiqué par le zoobenthos: 
tendance 1980-1998 


Claude LANG 
Conservation de la faune, Marquisat 1, CH-1025 St-Sulpice, Suisse. 
E-mail: claude.lang @sffn.vd.ch 


Trophic state of Lake Morat indicated by the zoobenthos: the 1980- 
1998 trend. - Total phosphorus concentrations have strongly decreased 
between 1982 and 1998 in the water of Lake Morat (Switzerland). In 
response to this improvement, the numbers of tubificids (Tubifex tubifex 
and Potamothrix hammoniensis mostly) decreased in the deepest area (40- 
45 m), but the other species (Limnodrilus profundicola and L. hoffmeisteri) 
present therein, remained scarce. In contrast at a depth of 20 m, chironomid 
larvae (Chironomus and Procladius) and the tubificid Potamothrix molda- 
viensis became more abundant in 1998. These changes indicated a slow 
improvement of benthic conditions but, in 1998, the zoobenthos was 
always characteristic for a eutrophic lake. 


Key-words: chironomid — eutrophication — indicator — lake — oligochaete - 
zoobenthos. 


INTRODUCTION 


Les concentrations en phosphore total dans l'eau du lac de Morat ont fortement 
diminué en réponse aux mesures d’assainissement prises dans le bassin versant, 
passant de 147 mg / m? en 1982 à 30 mg en 1998 (OPE, 1999). La baisse du phos- 
phore devrait entrainer celle de la production des algues planctoniques, donc une 
diminution de la sédimentation organique, et par conséquence une meilleure oxygé- 
nation des couches d’eau profondes (Sas, 1989). La production des algues n'a pas été 
mesurée mais les concentrations en oxygene ont augmente dans les eaux du fond 
pendant la stagnation estivale (OPE, 1999). Cette amélioration de l'oxygénation 
devrait faire sentir ses effets au niveau de la faune des sédiments profonds (faune 
benthique ou zoobenthos). Celle-ci se compose principalement (si on se limite a la 
macrofaune) d’oligochètes et de larves de chironomides (Diptera) dont les commu- 
nautés d'espèces varient en fonction de l’état du lac et peuvent ainsi servir d’indi- 
cateurs (Johnson et al., 1993). 

Des conditions oligotrophes (peu de phosphore et beaucoup d'oxygène) pré- 
valaient au début du siècle dans la plupart des grands lacs suisses (Sas, 1989). Les 
nombres d’individus appartenant a des especes d’oligochetes et de chironomides 


Manuscrit accepté le 26.01.2000 


234 CLAUDE LANG 


indicatrices de ce type de conditions ont diminué en réponse a l’augmentation du 
phosphore entre 1960 et 1980, tant dans le Léman que dans le lac de Neuchatel (Lang, 
1998, 1999). Lorsque ce nutriment a diminué, les nombres d’individus de ces espèces 
ont augmenté à nouveau dans ces deux lacs, montrant ainsi que l’état des sédiments 
profonds s’améliorait. Les espèces oligotrophes ne sont plus présentes dans la zone 
profonde du lac de Morat où les espèces caractéristiques des lacs eutrophes consti- 
tuent l’essentiel du zoobenthos entre 1980 et 1991 (Lang & Reymond, 1993). 

En effet, ce lac est eutrophe depuis longtemps. En 1825 déjà (Jaag, 1948), 
l’eau du lac de Morat était devenue rouge à cause de la prolifération d’une cyano- 
bactérie (Oscillatoria rubescens, appelée maintenant Planktothrix rubescens DC). 
Certains riverains, croyant que c'était le sang des soldats tués lors de la bataille de 
Morat en 1476 qui remontait à la surface, ont appelé ce phénomène Sang des 
Bourguignons. L’ apparition de cet organisme révèle que l’état d’un lac commence à 
se dégrader (Sas, 1989). En 1936, la rareté de l’oxygène dans les couches d’eau 
profondes pendant la stagnation estivale indique que les algues prolifèrent en surface; 
le Sang des Bourguignons est toujours abondant, mais pas de façon continue (Rivier, 
1936). La concentration en phosphore total qui est de 35 mg /m? en 1955, augmente 
jusqu’en 1982 avant de commencer à redescendre (Liechti, 1989). En 1972, le lac est 
qualifié d’eutrophe au vu de la composition chimique de ses sédiments (Davaud, 
1976). 

La présente étude cherche à évaluer et à interpréter la réponse du zoobenthos à 
la baisse des concentrations en phosphore entre 1982 et 1998 en termes d’amélio- 
ration de l’état biologique des sédiments profonds. En effet, le lac de Morat ne sera 
restauré que lorsque ses sédiments le seront puisqu'ils reçoivent, sous une forme ou 
une autre, tout ce qui est produit dans la colonne d’eau (Hakanson & Jansson, 1983). 
Si l’état des sédiments s’améliore, la densité des oligochètes devrait diminuer tandis 
que celle des larves de chironomides devrait augmenter (Wiederholm, 1980). De plus, 
la composition des espèces devrait se modifier au sein de ces deux groupes: les 
espèces très résistantes au manque d'oxygène devraient être remplacées peu à peu par 
celles qui prévalaient avant la phase d’eutrophisation intense (Lang, 1998). 


STATIONS ET MÉTHODES 


Le lac de Morat qui couvre une surface de 23 km?, est relativement peu 
profond (profondeurs moyenne et maximale: 23 m et 45 m respectivement) ce qui 
explique sa sensibilité vis-à-vis de l’augmentation des concentrations en phosphore 
(Liechti, 1989). Son principal affluent, la Broye, contribue pour 87 % à ses apports en 
phosphore (OPE, 1999); c’est donc surtout l’état du bassin versant de cet affluent qui 
détermine celui du lac. Les mesures d’assainissement prises ont entraîné la baisse 
dans l’eau du lac des concentrations en phosphore total de 62 mg / m? en 1987 à 31 
mg en 1996 (OPE, 1999). De ce fait, les concentrations moyennes en oxygène à 40 m 
de profondeur entre les mois de juillet et de novembre ont passé de 1.8 mg /1 en 1987 
à 3.6 mg / l en 1996; les valeurs extrêmes étant respectivement 1.5 à 2.9 mg et 2.1 à 
4.9 mg. Les années antérieures à 1987 et postérieures à 1996 ne sont pas incluses dans 
la comparaison parce que la fréquence des mesures d’oxygène n’y est suffisante ni 
pour calculer une concentration moyenne pendant l’été ni pour estimer la durée de la 


ZOOBENTHOS DU LAC DE MORAT 235 


période où l’oxygene est rare. Pour cette raison, il n'est pas possible de comparer 
directement l’évolution des concentrations en oxygène entre 1980 et 1998 a celle du 
zoobenthos. 

Entre 1980 et 1998, le zoobenthos du lac de Morat a été étudié dans 11 stations 
de prélévements localisées entre 10 m et 45 m de profondeur (Fig. 1). A cela 
s’ajoutent, en 1994, 42 stations réparties régulièrement sur l’ensemble du lac et, en 


Môtier 


Faoug I 
o) 


Fic. 1 


Localisation des onze stations de prélèvements visitées entre 1980 et 1998 dans le lac de Morat. 
Les flèches indiquent les points d’entrée et de sortie de la Broye, le principal affluent. 


1998, 15 stations distantes de 200 m les unes des autres, placées sur un transect qui va 
de la station 7 à la station 8. Dans chaque station, des carottes de sédiment de 16 cm? 
chacune ont été prélevées en nombre variable selon les années et les profondeurs 
(Tab. 1). Les prélèvements sont effectués au moyen d’un carottier, descendu depuis la 
surface à l’extrémité d’un cable, dans la plupart des stations, sauf dans les stations 1 et 
3, visitées en plongée par l’auteur en 1987 et en 1998. Seule la station 1 est située 
dans la zone littorale (10 m), toutes les autres le sont dans la zone profonde (20 m à 
45 m). 

En laboratoire, le sédiment contenu dans chaque carotte est tamisé (ouverture 
de maille: 0.2 mm). Le refus du tamis est fixé au formol 5 %. La faune benthique est 
ensuite séparée du sédiment, sous une loupe lumineuse à l’aide de pinces fines. Les 


236 CLAUDE LANG 


TABLEAU | 


Frequence et abondance (A) relatives (%) des especes de tubificides du lac de Morat en 
fonction de la profondeur, de la station et de l’année. Fréquences calculées seulement à partir 
des individus sexuellement matures. Fréquences 1980 pas comparables: + présence. Espece | 
caracteristique des lacs mésotrophes; especes 2 a 7 caractéristiques des lacs eutrophes (Milbrink 
1973): 


Profondeur (m) 


Especes 19: 10, 20.207220 20 203372402407 AO OISE 
1. Potamothrix moldaviensis 50 83 IM 

(Vejdovsky, Mrazek) 

2. Ilyodrilus templetoni 33 20 6 

(Southern) 

3. Limnodrilus claparedeanus 17 6 7 

Ratzel 

4. Limnodrilus hoffmeisteri 33 GTS 6 

Claparède 

5. Limnodrilus profundicola 17 17 10 + £25 19 31 + 12 

(Verrill) 

6. Potamothrix hammoniensis 83 + 25 ..6 100° ASE 87 IS CO- -S 
(Michaelsen) 

7. Tubifex tubifex (Miller) 337 920i a = oe 81 44 53 70 76 
Espece | (A) 1228019 2 

Espèces 3 à 5 (A) ID a ZITTI 7 200001 2 

Espèces 2, 6, 7 (A) 66,092. Sess) 7109 737,697 789293298 5999000 
Année $7 98 78571807, 8491 98" OA 808 AI EI See Oo RS 6 
Jour et mois 28.4 30-3) 877 B07 25:6925'6° 31.372167 30723.602 2031097238] 
Nbre carottes (16 cm2) 67212772105 1677 116 7167, 22 SRO IS 30 2021 
Nbre vers identifiés 40 48 24 121 125 59 56 402 169 306 143 134 170 174 


Stations (fig. 1) ] | 10) 2-5. 2-5), 9225) 3) Fig23)}6=9) 16-9) 6-9 Sl 


oligochetes, les larves de chironomides et de Chaoborus sont comptés et pesés 
(biomasse), apres passage sur du papier buvard pour enlever l’eau en exces. Tous les 
tubificidés (ou un sous-échantillon s'ils sont trop nombreux), dont le diamètre a l’état 
fixe dépasse 0.29 mm (Lang, 1998), sont montes entre lame et lamelle dans un milieu 
éclaircissant (Reymond, 1994) afin d’être identifié sous le microscope. Seuls les 
individus sexuellement matures des taxons présents dans le lac de Morat peuvent être 
identifies jusqu’au niveau de l’espece. Cependant, les immatures peuvent tout de 
méme étre classés dans des groupes d’especes dont la valeur indicatrice est analogue 
(Lang, 1998). Nous distinguons ainsi les especes caractéristiques des lacs oligo- 
trophes, mésotrophes ou eutrophes. La fréquence relative d’une espece s’exprime 
comme le pourcentage de carottes dans lesquelles cette espèce est présente. L’abon- 
dance relative d’une espèce se définit comme le nombre d’individus de cette espèce 
rapporté, sous forme de pourcentage, au nombre total d'individus identifiés dans une 
carotte ou un ensemble de carottes. 

De 1984 à 1998, les résultats sont analysés carotte par carotte, contrairement à 
1980, où les 4 carottes prises dans chacune des 8 stations visitées cette année-là sont 
cumulées. De plus, la biomasse n’a pas été mesurée en 1980. Pour ces raisons, les 


ZOOBENTHOS DU LAC DE MORAT 237 


résultats de 1980 ne peuvent pas étre inclus dans toutes les comparaisons effectuées 
entre les années. Les seules campagnes de prélevements vraiment comparables entre 
elles à tous les points de vue sont celles effectuées en 1984, 1991 et en 1998 a 20 m et 
40 m de profondeur. 

La biomasse du zoobenthos (g/m?) peut être calculée à partir des concen- 
trations en phosphore total (mg/m?) en utilisant une relation empirique établie dans 
d’autres lacs (Hanson & Peters, 1984): 


Log; biomasse = 0.708 log} phosphore + 0.092 


La concentration moyenne du phosphore, calculée sur les cinq années précédant le 
prélèvement du zoobenthos, est utilisée dans cette expression pour tenir compte du 
fait que le zoobenthos réagit souvent avec un temps de latence à la baisse du 
phosphore (Lang, 1998). Si la biomasse observée est proche de la biomasse calculée, 
cela signifie qu'il n’y a pas de décalage entre l’état trophique du sédiment et celui de 
l’eau. Ce type de relation empirique permet de replacer l’évolution trophique d’un lac 
particulier dans un contexte plus général et de mettre ainsi en évidence d'éventuelles 
anomalies (Lang, 1998). 


RÉSULTATS 


Dans le lac de Morat, la diversité du zoobenthos diminue avec la profondeur 
(Tab. 1) ce qui reflète la baisse des concentrations en oxygène (OPE, 1999). A 40 m 
et 45 m de fond, le zoobenthos se compose presque exclusivement de deux espèces de 
tubificidés, très résistantes au manque d'oxygène, dont la dominance numérique 
caractérise les lacs eutrophes: Potamothrix hammoniensis et Tubifex tubifex (Mil- 
brink, 1973). Le pourcentage de ces deux espèces dans les communautés d’oligo- 
chètes ne change guère à 40 m entre 1980 et 1998. A 20 m de profondeur, le pourcen- 
tage des espèces du genre Limnodrilus augmente par rapport à 40 m, mais reste assez 
constant entre 1980 et 1998. Notons en 1998 la présence à 20 m de Potamothrix 
moldaviensis, une espèce plutôt caractéristique des lacs mésotrophes (Milbrink, 
1973), qui est relativement abondante à 10 m, tant en 1987 qu’en 1998. Les cinq 
premières espèces du tableau 1 sont présentes dans le Léman jusqu’à 150 m de pro- 
fondeur au moins (Lang, 1998), ce qui montre que ce n’est pas ce facteur qui limite 
leur distribution verticale dans le lac de Morat, mais plutôt le manque d'oxygène. 

A 40 m de profondeur, la biomasse du zoobenthos ne change pas entre 1984 et 
1998 (Tab. 2), mais le nombre de tubificidés diminue significativement (test de 
Kruskal Wallis: P = 0.084 et P = 0.007 respectivement). À 20 m de fond, la biomasse 
du zoobenthos augmente entre 1984 et 1998. Cette augmentation est surtout le fait des 
larves de chironomides qui constituent en 1998 le 54 % de la biomasse. Seuls les 
genres Chironomus et Procladius, résistants au manque d’oxygène (Brinkhurst, 
1974), sont présents et leur abondance est la même (1500 individus/m?). Une ten- 
dance semblable s’observe en1998 à 10 m de profondeur: les chironomides forment le 
43 % de la biomasse, Chironomus et Procladius sont dominants. Mis à part 1998 (à 
10 m et 20 m), les chironomides ne représentent qu’une petite partie de la biomasse 
totale dans la zone profonde du lac de Morat qui se caractérise par la prédominance 
des tubificidés. Notons enfin la présence de quelques larves de Chaoborus à 40 m de 
profondeur. 


238 CLAUDE LANG 


TABLEAU 2 


Biomasse moyenne ( g / m2) du zoobenthos et nombre moyen d’oligochètes par m? en fonction 
de la profondeur et de l’année dans cing lacs: Morat, Léman, Neuchatel, Joux, Bret. Sources: 
cette étude, Lang 1998 et 1999, résultats non publiés. 


Biomasse 2) Nombre 
Profondeur ST 

Lac (m) Année nl) Moy. ESM Max. Moy. ESM Max. 
Morat 10 1987 6 13.5 3.1 24 30104 3016 39375 
1998 12 24.1 3.2 38 12604 1873 27500 

20 1980 16 - 3) = = 23750 à = 
1984 16 9.3 2.6 45 18085 2374 41250 
1991 16 7.8 1.5 19 11289 1341 25000 
1998 12 49.7 5.4 82 30677 3439 56875 
35 1994 42 43.2 8.5 356 49405 6074 245625 

40 1980 16 a = = 29375 = È 
1984 16 22.8 2.9 56 26875 3159 62500 
199] 16 19.9 3.9 66 21679 3408 55000 
1998 30 211 4.8 108 21541 5079 125625 
45 1985 20 33.1 6.5 119 21312 3212 53750 
1986 21 19.3 3.2 62 20833 2587 52500 
Léman 40 199] 192 24.0 1.6 219 24687 2111 274375 
1996 170 17.3 0.6 47 12092 585, 248105 
60 1994 159 13.5 0.6 38 20243 929 58750 
1999 159 11.6 0.7 57 12448 WS NHS 
Neuchatel 40 1992 171 10.2 0.5 35 9181 570 61250 
1997 175 9.4 0.5 32 8546 569 45625 
Joux 25 1998 450 52.1 1.5 186 39164 1001 230000 
Bret 20 1998 35 21:8 2.3 52 929 202 4375 


Leman 270 1998 25 4.0 0.8 19 5500 686 12500 


1) nbre de carottes de 16 cm? 
2) Moy. moyenne, ESM erreur standard de la moyenne, Max. valeur maximale. 
3) Résultats manquent. 


La campagne de prélévements effectuée en 1994 montre (Fig. 2 et 3) la 
répartition et la composition du zoobenthos dans l’ensemble la zone profonde du lac 
de Morat. Les valeurs de la biomasse du zoobenthos qui sont plus élevées dans la 
zone du lac influencée directement par les apports organiques de la Broye, le principal 
affluent (70 % des apports en eau, Liechti, 1989), diminuent en général au fur et a 
mesure que l’on s’en éloigne, tout comme les taux de sédimentation organique 
(Davaud, 1976). Tubifex tubifex et Potamothrix hammoniensis constituent l’essentiel 


ZOOBENTHOS DU LAC DE MORAT 239 


tant des tubificides (89 % des individus en moyenne) que du zoobenthos. Les autres 
especes de tubificidés du genre Limnodrilus se rencontrent surtout a la périphérie de 
la zone étudiée où la profondeur diminue. Les larves de chironomides ne sont 
présentes que dans 6 des 42 stations visitées et, en nombre d’individus (un individu 
seulement dans chacune des 6 stations), elles ne représentent que le 0.2 % du nombre 
des tubificidés. Cependant, comme les prélèvements ont été effectués le 14 et le 21 
juin, donc à la fin de la période d’émergence des chironomides, l’abondance des 
larves est probablement sous-estimée. Des prélèvements effectués en avril seraient 
plus représentatifs à cet égard. 


569 570 571 572 573 574 575 576 


FIG. 2 


Répartition géographique de la biomasse du zoobenthos (g /m?) dans le lac de Morat en 1994. 
Les coordonnées géographiques de la carte nationale de la Suisse sont indiquées en x et en y. 


La biomasse moyenne du zoobenthos est de 43.2 g / m? dans les 42 stations 
visitées en 1994 (Fig. 2); la biomasse médiane qui ne prend pas en compte les stations 
très influencées par les apports de la Broye, est de 32.2 g / m?. Ces deux valeurs 
observées sont bien supérieures à 16.5 g / m?, la biomasse calculée à partir de la 
moyenne (39 mg / m?) des concentrations en phosphore dans l’eau entre 1989 et 1993 
(voir Stations et méthodes). Par contre, la biomasse médiane observée en 1994 est très 
proche de 35.2 g / m?, la biomasse calculée à partir de la moyenne des concentrations 
en phosphore mesurées entre 1982 et 1986 (113.4 mg / m?). Cette différence montre 
que la biomasse du zoobenthos réagit lentement à la baisse du phosphore si bien que 


240 CLAUDE LANG 


199.5 
10074722100210033285 
199.0 - D a Oo o D DO 
33. 66 710071007100310053 
198.5 - o o o o o o o 
73 64 100 100 100 100 100 100 
198.0 - [n] [m] je} D je} O D D 
| 100 100 100 100 100 67 100 100 
197.5 D D D Oo o D D D 
62 94 100 100 8 71100 
197.0- (a) D D Oo [=] [n] [a] 
Broye 
100 100 91 70 100 100 
196.5 + o o o o o o 
196.0 | U T T T T 
569 570 571 572 57S 574 575 576 


FIG. 3 


Répartition géographique du pourcentage de Tubifex tubifex et de Potamothrix hammoniensis 
dans les communautes de tubificidés du lac de Morat en 1994. 


les sédiments restent capables en 1994 de nourrir des populations benthiques qui 
correspondent encore au niveau de productivité observé dix ans plus tot dans l’eau du 
lac. 


DISCUSSION 


Le lac de Morat est un des premiers lacs suisses dont l’état s’est dégradé sous 
l’influence des activités humaines (Jaag, 1948). La prolifération du Sang des 
Bourguignons en 1825 indique le début de cette dégradation. Une intensification des 
activités agricoles dans le bassin versant de la Broye, le principal affluent du lac, 
pourrait en avoir été la cause, comme c’est le cas depuis le moyen age déja pour de 
nombreux lacs d’Europe (Anderson, 1995). Signalons par exemple la culture du tabac 
qui s’est développée entre 1719 et 1819 dans la plaine de la Broye (Chuard & 
Dessemontet, 1972 ). De même, la production de blé a augmenté de 38 % entre 1707- 
1709 et 1787-1789 dans les balliages de Payerne et d’Avenches qui englobaient la 
partie inferieure du bassin versant (Chevallaz, 1949). Cet auteur fait également 
remarquer que l’agriculture vaudoise s’est fortement développée a partir de 1803 
grace à la suppression des usages et des structures contraignants de l’ Ancien Régime. 


ZOOBENTHOS DU LAC DE MORAT 241 


Cette evolution rapide des pratiques agricoles s’effectue dans un contexte 
météorologique particulier. Les années 1816 et 1817 se caractérisent en effet par des 
pluies et des inondations exceptionnelles (Pfister, 1985). Celles-ci résultent du 
bouleversement climatique engendré par la présence, dans la haute atmosphere, des 
poussières provenant de l’éruption en 1815 du volcan indonésien Tambora (Whyte, 
1995). Ces pluies, tombant sur les sols du bassin versant du lac de Morat, cultivés de 
façon plus intensive, en ont probablement lessivé les nutriments, créant ainsi les 
conditions qui ont provoqué en 1825 la prolifération du Sang des Bourguignons. Cette 
longue histoire de dégradation (plus de 150 ans) laisse supposer que la restauration 
biologique du lac de Morat prendra plus de temps que celle du Léman ou du lac de 
Neuchatel ou la phase de détérioration n’a duré qu’une trentaine d’années (Lang, 
1998, 1999). 

Malgré la baisse spectaculaire des concentrations en phosphore dans l’eau du 
lac de Morat, la composition du phytoplancton et son abondance, ainsi que celles du 
périphyton épilithique, restent caractéristiques d’un lac eutrophe en 1988 et en 1989 
(Reymond & Straub, 1993). De même, le zoobenthos ne se modifie pas de façon 
fondamentale entre 1980 et 1998, tout au moins dans la zone profonde, seule étudiée 
en détails. Les deux espèces de tubificidés qui restent les plus abondantes entre 1980 
et 1998, Tubifex tubifex et Potamothrix hammoniensis, étaient déjà les seules 
présentes en 1935 dans la zone la plus profonde du lac (Rivier, 1936). A cette époque, 
elles colonisaient en petit nombre le fond du lac en compagnie des larves de 
Chaoborus qui étaient relativement abondantes. Ce genre de faune se rencontre dans 
des lacs très eutrophes, où le manque d’oxygene persiste pendant des mois. Dans ce 
type de situation, l'abondance des tubificidés qui vivent constamment en contact avec 
le fond du lac, diminue davantage que celle des Chaoborus qui peuvent migrer 
chaque nuit vers la surface et les couches oxygénées (Brinkhurst, 1974). C’est la 
situation observée en 1998 dans le lac de Bret à 20 m de profondeur où Chaoborus 
constitue le 98 % de la biomasse (Tab. 2). Lorsque le lac est profond, les Chaoborus 
sont absents: dans le Léman par exemple, seuls les tubificidés sont présents entre 270 
m et 300 m de fond et leur biomasse est faible à cause du manque d’oxgene qui 
caractérise cette zone (Lang, 1998). En 1972 comme en 1936, les sédiments profonds 
du lac de Morat semblaient presque dépourvus de faune, mis à part les Chaoborus et 
quelques tubificidés (Davaud, 1976). Entre 1980 et 1998, les tubificidés sont devenus 
abondants et les Chaoborus rares, ce qui tendrait à montrer que l’état biologique des 
sédiments s’est amélioré par rapport à la situation précédente. Toutefois, comme 
celle-ci n’est pas décrite de façon quantitative dans les travaux de Rivier et de 
Davaud, l'interprétation de la tendance peut être mise en doute. 

Lorsque l’état d’un lac s'améliore après un épisode d’eutrophisation, la densité 
des larves de chironomides augmente tandis que celle des oligochètes diminue 
(Wiederholm, 1980). Cette tendance s’observe en 1998 dans le lac de Morat: à 10 m 
et à 20 m de profondeur en ce qui concerne les chironomides, à 40 m en ce qui 
concerne les tubificidés. Remarquons toutefois que l’abondance moyenne des oligo- 
chètes en 1998 à 40 m est plus du double de celles observées à la même profondeur 
dans le lac de Neuchâtel en 1992 et 1997 (Tab. 2); les valeurs du Léman à 40 m et à 
60 m de profondeur se situent entre ces deux extrêmes. Cette séquence des abon- 
dances — Morat, Léman, Neuchâtel — reflète la décroissance des concentrations en 


242 CLAUDE LANG 


phosphore entre ces trois lacs. Cependant, il existe des exceptions à cette règle. Dans 
le lac de Joux par exemple, malgré la baisse du phosphore, la biomasse du zoobenthos 
et l’abondance des tubificidés restent élevées à cause de la présence presque continue 
du Sang des Bourguignons qui sédimente en masse sur le fond du lac entre 1980 et 
1998. Une fois installé dans un lac, cet organisme peut en effet s’y maintenir malgré 
les mesures d’assainissement prises (Sas, 1989). Son retour en force dans le lac de 
Morat n’est d'ailleurs pas exclu. 

L’amélioration constatée dans le lac de Morat est plus marquée à faible qu’à 
grande profondeur. De ce fait, des prélèvements de zoobenthos concentrés sur les 
bords de la zone profonde (entre 15 m et 30 m) auraient probablement mieux mis en 
évidence la restauration de l’état des sédiments que le programme suivi. Une autre 
strategie possible consiste a augmenter l’intervalle de temps entre les campagnes de 
prélevements de maniere a ce que le zoobenthos puisse répondre de facon plus claire 
à la baisse du phosphore. Par exemple, la campagne de 1994 qui couvre l’ensemble de 
la zone profonde (Fig. 2 et 3), pourrait étre refaite en 2004, si possible en avril avant 
l’émergence des chironomides. Si ces dix ans d’amélioration chimique ont eu un effet 
positif au niveau du sédiment, les larves de chironomides devraient devenir plus 
nombreuses et recoloniser l’ensemble de la zone profonde, la baisse de l’abondance 
des tubificidés devrait se poursuivre et la dominance de Tubifex tubifex et Potamo- 
thrix hammoniensis devrait diminuer au profit d’autres espèces moins résistantes au 
manque d'oxygène. La colonisation par Potamothrix moldaviensis, espèce indicatrice 
de conditions mésotrophes, des sédiments situés à 45 m de profondeur montrerait que 
le lac de Morat est vraiment en voie de restauration. 

Si la baisse du phosphore se poursuit, le retour des espèces indicatrices de 
conditions oligotrophes, comme dans le Léman et le lac de Neuchâtel (Lang, 1998, 
1999), n’est pas exclu à long terme. L’exemple du lac Esrom montre que celles-ci 
peuvent en effet rester présentes à très faible profondeur (entre 0.5 m et 1.5 m) dans le 
littoral d’un lac eutrophe, comme c’est le cas du lumbriculidé Srylodrilus heringianus 
(Dall et al., 1990). Signalons à ce propos qu’en 1905 Spirosperma ferox et Psammo- 
ryctides barbatus, deux espèces de tubificidés indicatrices de conditions oligo- 
mésotrophes (Milbrink, 1973), étaient présentes à 4 m de profondeur dans le lac de 
Morat (Piguet, 1906). Si elles ont subsisté à très faible profondeur comme c’est 
probable, elles pourraient graduellement recoloniser le lac au fur et à mesure des 
progrès de la restauration. 


REMERCIEMENTS 


L’aide d'Olivier Reymond en laboratoire et celle de Raymond Ducret sur le 
terrain m'ont permis de mener à bien ce travail. Les remarques du comité de lecture 
ont contribué à améliorer cet article. 


BIBLIOGRAPHIE 

ANDERSON, N. J. 1995. Naturally eutrophic lakes: reality, myth or myopia? Trends in Ecology 
and Evolution 10: 137-138. 

BRINKHURST, R. O. 1974. The benthos of lakes. The Macmillan Press, London, 190 pp. 


CHEVALLAZ, G. A. 1949. Aspects de l'agriculture vaudoise à la fin de l’Ancien Régime. F. 
Rouge, Lausanne, 272 pp. 


ZOOBENTHOS DU LAC DE MORAT 243 


CHUARD, J. P. & DESSEMONTET, O. 1972. Le 250° anniversaire de la culture du tabac en pays 
romand: 1719-1969. Imprimeries Réunies, Lausanne, 73 pp. 

DALL, P. C., LINDEGAARD, C. & JONASSON, P. M. 1990. In-lake variations in the compositions 
of zoobenthos in the littoral of Lake Esrom, Denmark. Verhandlungen der internatio- 
nale Vereinigung fiir theoretische und angewandte Limnologie 24: 613-620. 

DAVAUD, E. 1976. Contribution a l’étude géochimique et sédimentologique de dépôts lacustres 
récents (lac de Morat, Suisse). Thése, Genéve, 129 pp. 

HAKANSON, L. & JANSSON, M. 1983. Principles of lake sedimentology. Springer-Verlag, Berlin, 
316 pp. 

HANSON, J. M. & PETERS, R. H. 1984. Empirical prediction of crustacean zooplankton biomass 
and profundal macrobenthos biomass in lakes. Canadian Journal of Fisheries and 
Aquatic Sciences 41: 439-445. 

JAAG, O. 1948. Die neuere Entwicklungen und der heutige Zustand der Schweizer Seen. 
Verhandlungen der internationale Vereinigung fiir theoretische und angewandte Lim- 
nologie 10: 192-209. 

JOHNSON, R. K., WIEDERHOLM, T. & ROSENBERG, D. M. 1993. Freshwater biomonitoring using 
individual organisms, populations, and species assemblages of benthic macroinver- 
tebrates. In: Rosenberg, D. M. & Resh, V. H. (eds). Freshwater biomonitoring and 
benthic macroinvertebrates. Chapman & Hall. New York, 488 pp. 

LANG, C. 1998. Using oligochaetes to monitor the decrease of eutrophication: the 1982 - 1996 
trend in Lake Geneva. Archiv fiir Hydrobiologie 141: 447-458. 

LANG, C. 1999. Contrasting responses of oligochaetes (Annelida) and Chironomids (Diptera) to 
the abatement of eutrophication in Lake Neuchatel. Aquatic Sciences 61: 206-214. 

LANG, C. & REYMOND, O. 1993. Eutrophisation du lac de Morat indiquée par les communautes 
d’oligochètes: tendance 1980 - 1991. Revue Suisse de Zoologie 100 (1): 11-18. 


LIECHTI, P. 1989. L’état du lac de Morat. Bulletin de l'OFEFP, Berne. 2/89: 32-36. 

MILBRINK, G. 1973. On the use of indicator communities of tubificidae and some lumbriculidae 
in the assessment of water pollution in Swedish lakes. Zoon 1: 125-139. 

OPE 1999. Office de Protection de l'Environnement du canton de Fribourg. Rapport interne. 
Fribourg. 

PIGUET, E. 1906. Oligochètes de la Suisse française. Revue Suisse de Zoologie 14: 398-403. 

PFISTER, Chr. 1985. Klimageschichte der Schweiz. Verlag Paul Haupt Bern und Stuttgart. 184 
PP- 

REYMOND, O. 1994. Préparations microscopiques permanentes d’oligochètes: une methode 
simple. Bulletin de la Societe Vaudoise des Sciences Naturelles 83: 1-3. 

REYMOND, O. & STRAUB, F. 1993. Phytoplancton du lac de Morat en 1988-1989: comparaison 
avec les données anterieures. Bulletin de la Société Neuchäteloise des Sciences Natu- 
relles 116: 55-64. 

RIVIER, O. 1936. Recherches hydrobiologiques sur le lac de Morat. Bulletin de la Société 
Neuchäteloise des Sciences Naturelles 61: 125-180. 

SAs, H. 1989. Lake restoration by reduction of nutrient loading. Academia Verlag Richarz 
GmbH. St. Augustin. 497 pp. 

WHYTE, I. D. 1995. Climatic change and human society. Arnold London 217 pp. 

WIEDERHOLM, T. 1980. Use of benthos in lake monitoring. Journal of Water Pollution and 
Control Federation 52: 537-547. 


REVUE SUISSE DE ZOOLOGIE 107 (2): 245-257; juin 2000 


A new species of Astyanax (Characiformes, Characidae) 
from Uruguay river basin in Argentina, with remarks 
on hook presence in Characidae 


Maria de las Mercedes AZPELICUETA! and José O. GARCIA? 

| Dep. Cientifico Zoologia de Vertebrados, Fac. de Ciencias Naturales y Museo de 
La Plata, Paseo del Bosque, 1900 La Plata, Argentina. 

? Fac. de Cs. Exactas y Naturales, Universidad Nacional de Misiones, 
Rivadavia 2370, 3300 Posadas, Argentina. 


A new species of Astyanax (Characiformes, Characidae) from Uruguay 
river basin in Argentina, with remarks on hook presence in Chara- 
cidae. - In the present paper the new species Astyanax ojiara sp. n. is 
described, from the headwaters of Yaboti river, an affluent of Uruguay river 
in the province of Misiones, Argentinean northeast. A combination of 
characters differentiates the new species from other congeners: one 
heptacuspid maxillary teeth; teeth of inner premaxillary row gently expan- 
ded distally; 7-9 dentary teeth decreasing in size anteroposteriorly; males 
with hooks in all fins; large males with pelvic axillary scale bearing | to 8 
hooks; 36-38 perforated scales in lateral line; 20-23 branched anal-fin rays. 
Number of scales in the lateral line and branched anal-fin rays, humeral 
spot, and low body of Astyanax ojiara sp. n. resemble those of A. eigen- 
manniorum, but maxillary, premaxillary and dentary teeth differ in both 
species. The disposition of the dentary teeth is also similar in A. taeniatus 
and A. giton but A. ojiara sp. n. has one heptacuspid maxillary tooth. 


Key-words: Characiformes - Characidae - Astvanax - new species - 
Uruguay river. 


INTRODUCTION 


In the last years, the genus Astyanax was not revised and an accurate definition 
of the genus is still pendant. Eigenmann (1921, 1927) analised the genus and, sub- 
sequently, many authors followed his results. Géry (1977) provided an arrangement of 
the species in groups, and identification keys. Recently, Zanata (1997) erected a new 
genus - Jupiaba - for many species previously known as Astyanax or Deuterodon. 
Jupiaba is defined by the presence of a diagnostic pelvic bone, with unique disposition 
of muscles. The taxonomic status of some species of the genus Astyanax is not comple- 
tely clear and the interrelationships of the species remain unresolved. A detailed 
revision of the remaining nominal species of Astyanax is necessary. 


Manuscript accepted 27.12.1999 


246 MARIA DE LAS MERCEDES AZPELICUETA & JOSE O. GARCIA 


In the rio de la Plata basin, about twenty nominal species of Astyanax have been 
recorded, although the records of some species are doubtful. In the southwest of the 
Brazilian shield, the main course of the rio Uruguay flows from East to West, turning to 
the southwest and ending in the rio de la Plata. Some affluents of the river originate 
from the Sierra de Misiones, which reach about 800 m a.s.l., and presently constitute an 
effective barrier between waters of the Uruguay and Paranä rivers, in northeastern 
Argentina. The arroyo Yaboti comes from the highest area of Sierra de Misiones; in one 
of its headwaters, the arroyo Benitez, a new species of Astyanax was collected in the 
vicinity of the city of San Pedro. The description of the new species is presented in this 


paper. 


MATERIAL AND METHODS 


The specimens examined in this study were cleared and counterstained (C&S) 
following Taylor & Van Dyke (1985). Measurements are straight distances taken with 
calliper to nearest 0.1 mm. Material is deposited in the Field Museum of Natural 
History, Chicago (FMNH); Muséum d’histoire naturelle de Geneve, Geneve (MHNG); 
Facultad de Ciencias Naturales y Museo de La Plata, La Plata (MLP); Museu de 
Zoologia da Universidade de Sao Paulo, Sao Paulo (MZUSP). 

Comparative material (SL in mm). Astyanax eigenmanniorum (Cope, 1876): 
MLP 5202, 5 ex., 56.5-68.5, Argentina, Cordoba, rio Primero frente a Capilla de los 
Remedios; MLP 9160, 6 ex., 36.8-80.2, Argentina, Buenos Aires, Los Talas. Cleared 
and stained material: Personal collection of MA. Astyanax abramis (Jenyns, 1842): 2 
ex., 74.5-92.0, Argentina, Buenos Aires, rio de la Plata en Punta Lara; 2 ex., 80.6-98.8, 
Argentina, Misiones, rio Piray-Mini. Astyanax alburnus (Hensel, 1870): 5 ex., 40.5- 
47.2 , Uruguay, rio Yaguarön. Astyanax alleni (Eigenmann and McAtee, 1907): 2 ex., 
62.4-72.9, Argentina, Corrientes, rio Riachuelo; 1 ex., 66.0, same collecting data. 
Astyanax cf. asuncionensis Géry, 1972: 2 ex., 28.0-37.6, Argentina, Santa Fe, Isla Los 
Sapos; 2 ex., 80.4-92.7, Argentina, Misiones, rio Uruguay en San Isidro. Astyanax 
eigenmanniorum: 2 ex., 17.7-33.0, Argentina, Buenos Aires, desembocadura del rio 
Colorado; 2 ex., 28.0-30.5, Argentina, Buenos Aires, Laguna de Gomez; 1 ex., 60.3, 
Argentina, Buenos Aires, rio de la Plata; 4 ex., 51.5-82.1, Argentina, Misiones, arroyo 
Piray-Mini; 1 ex., 45.0, Brasil, Rio Grande do Sul, Viamäo, agude Charolés; Astyanax 
cf. fasciatus (Cuvier, 1819): 2 ex., 91.0-106.5, Argentina, Misiones, rio Uruguay en San 
Isidro. Astyanax ojiara sp. n.: 3 females, 46.2-63.0; 7 males, 37.8-58.0, Argentina, 
Misiones, arroyo Benitez. 


TAXONOMY 
Astyanax ojiara Sp. n. Figs 1-13, table 1 


Holotype (Fig. 1). MLP 9470, male, 50.5 mm SL, Argentina, province of Misiones, 
arroyo Benitez, headwaters of rio Yaboty, an affluent of rio Uruguay, coll. J. O. Garcia, May 
1983. 

Paratypes. (Collecting data as holotype). MLP 9471, 2 males, 38.0-45.0 mm. MLP 9472, 
12 females, 39.8-72.0 mm. MHNG 2605.67, 10 males (measured); MHNG 2606.35, 3 females, 
44.0-53.5 mm, (not measured). FMNH 98319, 5 ex. MZUSP 40255, 5 ex. 


A NEW SPECIES OF ASTYANAX FROM ARGENTINA 247 


DIAGNOSIS 


The species is distinguished by a combination of characters: one maxillary teeth 
with seven small cusps; teeth of inner premaxillary row gently expanded distally; 7-9 
dentary teeth decreasing in size anteroposteriorly; large males with hooks ın all fins; 
large males with | to 8 hooks in the pelvic axillary scale; 36-38 perforated scales in the 
lateral series; and iv-v,20-23 anal-fin rays. Also, presence of humeral spot vertically 
elongated, a second faint humeral spot, caudal spot continued on middle caudal rays, 
and low body depth help to differentiate the new species. 


DESCRIPTION 


Morphometrics of holotype and 24 paratypes are presented in table 1. Astyanax 
with low body (Fig. 1), maximum body depth at dorsal-fin origin. Dorsal profile of 
body slightly convex from snout to posterior tip of supraoccipital process, angled 
behind supraoccipital, strongly marked in medium sized and large females. Dorsal 
profile of body gently curved from this point to origin of dorsal fin; in large females, 
that portion almost straight; slanted ventrally from dorsal-fin origin to caudal peduncle. 
Dorsal profile of caudal peduncle straight; ventral profile slightly convex or straight. 
Ventral profile of body slightly curved from tip of snout to pelvic-fin origin, straight 
between this point and anal-fin origin, and slanted dorsally to caudal peduncle. Body 
rounded between pectoral and pelvic fins. Body laterally compressed between pelvic 
and anal fins. 

Dorsal-fin origin nearer tip of snout than base of caudal-fin rays, equally distant 
in some small specimens. Pelvic-fin origin anterior to vertical through dorsal fin-origin. 
Adipose fin anterior to bases of last branched anal-fin rays. Tip of pectoral fin falling 
near, sometimes surpassing, pelvic-fin origin; in large females, tip of pectoral fin far 
from that origin. Tip of pelvic fin reaching anal-fin origin in males; in females, pelvic- 
fin tip far from that origin, even in many small specimens. 

Dorsal fin iii, 9; posterior margin of dorsal fin slightly rounded, first branched 
dorsal-fin ray longest. In males, all dorsal-fin rays, excluded last one and unbranched 
rays, with slender hooks, directed outward, most of them curved ventrally, more 
abundant on posterior branch of each ray (Fig. 2), usually, one pair on each segment. 
Few hooks developed on ray tips of small specimens. 

Anal-fin iv-v, 20-23 (holotype 21). Posterior margin almost straight in males; in 
females, first five branched rays produced forming a small lobe. Anal fin of all males 
bearing hooks directed posteriorly and outward, slightly curved dorsally. Large 
specimens with hooks on last branched anal-fin ray and also on largest unbranched ray. 
Hooks placed on all branches of the ray (Fig. 3), sometimes two or three pairs on each 
segment. 

Caudal fin bearing 10-12 dorsal and 9-10 ventral procurrent rays; one 
unbranched and 9 branched principal rays in upper lobe; 8 branched and 1 unbranched 
principal rays in lower lobe. Usually, caudal lobes similar in size; sometimes, lower one 
slightly longer. Few hooks occurring on distal tips of middle caudal-fin rays in males 
(Fig. 4). In some specimens, few hooks scattered on all fin rays, included principal 
unbranched one. Hooks directed outward, most of them on lower lobe curved dorsally 


248 MARIA DE LAS MERCEDES AZPELICUETA & JOSÉ O. GARCIA 


FIG. 1 


Astyanax ojiara new species, holotype, MLP 9470, male, SL 50.5 mm, Argentina, province of 
Misiones, arroyo Benitez, headwaters of rio Yaboty, an affluent of rio Uruguay. 


and those ones of upper lobe curved ventrally. Some small specimens with 2 or 3 pairs 
of hooks only; one 43.0 mm SL male without hooks. 

Pectoral-fin 1, 10-13 (holotype i, 12), rays bearing hooks of different size in 
males. In medium sized and large specimens hooks developed on first unbranched ray 
and until 9 branched rays. Sometimes more than one pair of hooks on each segment, 
some of them directed dorsally and other ones ventrally, most of them directed inward 
(Fig. 5). Very few hooks developed in small specimens. 

Pelvic fin i, 7; hooks developed on branched rays, occasionally on first 
unbranched ray also in large specimens (Fig. 6). One or two pairs of hooks on each 
segment, in different branches of ray, most of them directed inward, slightly curved 
anteriorly. Usually, one ray with scarce hooks in small specimens. 

Head length moderate, mouth terminal, horizontal; snout short. Lower jaw 
slightly longer. Premaxilla with short ascending process, bearing two series of teeth. 
Usually, 3 pentacuspid or tricuspid teeth in outer series, sometimes two or four teeth, 
central cusp larger. Inner series of premaxillary teeth consisting of 5 teeth, gently 
expanded distally, slightly compressed at distal tips (Fig. 8). Symphysial tooth narrower 
and deeper, with 4 or 6 cusps. Second tooth widest, with 7 cusps. Third and fourth teeth 
with 6-7 cusps. Fifth tooth smaller, with 5 cusps. In all teeth, central cusp slightly larger 
than remaining ones. Maxilla short, scarcely lobed posteriorly, almost reaching or 
scarcely surpassing vertical through anterior orbital margin. One compressed maxillary 
tooth, with seven small cusps (Fig. 9). Few specimens (about 8 %) with two maxillary 
teeth, one of them heptacuspid. Dentary bearing 7 to 9 (usually 8) teeth with broad 
bases, decreasing in size anteroposteriorly. Symphysial tooth narrower, with 5-6 cusps. 
Second tooth widest. Second, third and fourth teeth with 5-7 cusps. Fifth tooth with 3-5 


A NEW SPECIES OF ASTYANAX FROM ARGENTINA 249 


Fics 2-4 


Astyanax ojiara, left view. 2: dorsal fin, SL 44.5 mm, scale= 1 mm; 3: detail of hooks on third 
branched anal-fin ray, SL 39.6 mm, scale= 0.5 mm; 4: detail of hooks on middle caudal-fin rays, 
SL 39.6 mm, scale= 1 mm. 


250 MARIA DE LAS MERCEDES AZPELICUETA & JOSE O. GARCIA 


cusps, sixth and seventh teeth with 1 to 3 cusps; eighth tooth - and nineth one when 
present - always conic (Fig. 10). 

Eye small, interorbital wide. Six infraorbitals well developed; third infraorbital 
almost contacting sensory tube of preopercle. Anterior fontanelle triangular, widening 
posteriorly; posterior fontanel long, extending onto supraoccipital process base. 

Scales cycloid, crenate. Lateral series with 36-38 perforated scales (holotype 
with 37, one specimen with 39). Lateral line running on lower half of caudal peduncle, 
ending in a long tube without lamina, between caudal rays. Five scales between dorsal- 
fin origin and lateral line; 5 between lateral line and ventral-fin origin. Thirteen or 
fourteen scales around caudal peduncle. Eleven or twelve scales forming a regular row 
between supraoccipital process and dorsal-fin origin; sometimes, 14 scales in an 
irregular row. Ten to twelve (until 14) rectangular scales placed on anal-fin base, 
covering all unbranched and twelve branched anal-fin rays. Scales placed on basal fifth 
of caudal lobes. A narrow axillary scale present dorsal to pelvic-fin insertion, oval, as 
long as one third of pelvic fin sometimes. Axillary scale bearing two hooks in its 
posterior inner area in medium and large males; although, a high number of hooks 
developed (8 in largest male, 56 mm SL, Figs 11-13). Young males with a ridge in that 
area of scale. 

In ten cleared and stained specimens, first arch bearing 17-19 gill-rakers: 2 on 
hypobranchial, 8 on ceratobranchial, 1 on cartilage, and 6 to 8 on epibranchial. 
Vertebral counts including Weberian apparatus and CUI+PUI as one element: 32 (1 
sp.), 33 (2 sp.), and 34 (7 sp.). 

Coloration of alcohol preserved specimens: Background light brown, dorsal 
region of flanks and head darker. A humeral spot well developed, dorsoventrally 
expanded, at level of pectoral-fin origin. A second lateral spot faint and smaller, usually 
rounded. A dark lateral band with different intensity in coloration and width crossing 
flanks. A deep and very narrow line of chromatophores runnig from humeral spot to 
base of middle caudal-fin rays. Most specimens with dark chromatophores on inner 
opercular surface forming a spot. 

Dark chromatophores on distal margin of anal fin, forming a band; two or three 
first dorsal rays with black chromatophores; middle caudal-fin rays black, also tips of 
caudal-fin rays with dark chromatophores; pectoral and pelvic fins hyaline. 


ETYMOLOGY 


The specific epithet is the name of a spirit, protector of the fresh waters, in the 
tupi-guarani language. 


DISCUSSION 


Eigenmann (1921, 1927) separated the species of the genus Astyanax into three 
subgenera that were maintained by many authors. Astyanax ojiara has low body depth 
and a complete series of scales in the predorsal area. Following Géry (1977), A. ojiara 
is placed in the group with 32-41 scales in the lateral line. Among those species, the 
number of anal-fin rays of A. ojiara agrees with that of A. fasciatus group. Like Géry 
(1977) pointed out, the other group is defined by the presence of 17-24 anal-fin rays, 


A NEW SPECIES OF ASTYANAX FROM ARGENTINA 25] 


GE 


= 


er 9 
Za 


N 


EI 


es Ss Ee 
7 
a 


Fics 5-7 
Astyanax ojiara, dorsal view, SL 45.0 mm. 5: left pectoral fin, scale= 1 mm; 6: detail of hooks on 
second branched pelvic-fin ray, scale= 0.5 mm. 7: A. eigenmanniorum, dorsal view, SL 45.0 mm, 
detail of the same ray, scale= 0.5 mm. 


252 MARIA DE LAS MERCEDES AZPELICUETA & JOSE O. GARCIA 


except A. eigenmanniorum. The number of perforated scales in the lateral line and the 
number of anal-fin rays of A. ojiara ressemble those of A. eigenmanniorum. 
Nonetheless, the number and shape of teeth greatly differ. Astyanax eigenmanniorum 
has 4-5 teeth in the outer series of premaxilla and 5 teeth in the inner one, with the 
central cusp notably larger; the maxillary tooth has 3 to 5 cusps (Azpelicueta, 1979); all 
those teeth always have broad bases. The dentary bears 4 large teeth, a median one, and 
a series with 5 to 7 very small teeth. In comparison, the maxilla of A. eigenmanniorum 
is narrow and long, always reaching the anterior third of the eye; the premaxilla has a 
longer dorsal process. Astyanax eigenmanniorum, as described by Cope (1876), has 
pectoral fins that reach ventral-fin origin, a character always present in the Brazilian 
specimens examined; the length of pelvic fins and the preanal distance have similar 
values that those of males of A. ojiara. The pelvic-fin hooks of A. eigenmanniorum are 
strong and regularly placed (Figs 6, 7); the anal-fin hooks develop on the first ten rays 
whereas they are smaller, more numerous and are present on many anal-fin rays of A. 
ojiara. In A. eigenmanniorum, the eye, the postorbital length, and the interorbital 
distance are larger. 

From other species of the A. eigenmanniorum group, A. ojiara is distinguished 
by one maxillary heptacuspid tooth, and the hooks developed in all fins of males. 
Astyanax taeniatus (Jenyns, 1842) and A. giton Eigenmann, 1908 have dentary teeth 
decreasing in size anteroposteriorly. Lucena & Lucena (1992) discussed the presence of 
that feature as a character that evolved in different species of the genera Deuterodon 
and Astyanax. 

Different measurement relations in both sexes of A. ojiara are the length of 
pelvic fin, the distances between last branched anal-fin ray-hypural joint, snout tip-anal- 
fin origin, and origins of pelvic and anal fins (Table 1). Two thirds of the sample 
examined were males. 


Presence of hooks in Characidae 


Sexual dimorphism appears in many characids; hooks develop in different fins 
of males, excluding the genus Cheirodon in which female pelvic fins bear hooks 
(Malabarba, 1998, fig. 16). One of the attributes of secondary sexual characteristics is 
its appearance during breeding season, followed by regression. Nonetheless, the speci- 
mens of A. ojiara examined were collected in May -the end of fall in southern 
hemisphere-, and neither males nor females were mature. Therefore, hooks are always 
present in the new species; this fact has been observed in the genera Cheirodon and 
Hyphessobrycon as well. 

In several subfamilies of Characidae such as Iguanodectinae, Stethaprioninae, 
Glandulocaudinae or Cheirodontinae and in different lineages within the family, males 
usually show hooks on anal and pelvic fins, but sometimes they occur on caudal or 
dorsal fins. Within the Cheirodontinae, some species as Serrapinus calliurus bears 
hooks in anal fin while many other ones have hooks on anal and pelvic fins. Among 
them, C. pisciculus, C. terrabae, C. dialepturus, C. affinis, C. gorgonae (Fink & 
Weitzman, 1974), C. ortegai (Vari & Géry, 1980), and C. interruptus (pers. obs.), 
Serrapinus kriegi and S. microdon (Uj, 1987; pers. obs.); and Heterocheirodon yatay 


A NEW SPECIES OF ASTYANAX FROM ARGENTINA 253 


Fics 8-13 


Fics 8-10: Astyanax ojiara, lingual view of left teeth, SL 48.0 mm, scale= 0.5 mm. 8: inner row 
of premaxillary teeth, with tips gently expanded distally; 9: maxillary tooth with seven small 
cusps; 10: eight dentary teeth, decreasing in size anteroposteriorly. Fics 11-13. Astyanax ojiara, 
left pectoral axillary scale of males, SL 42.0 mm; 11: lateroventral view, scale= 0.5 mm; 12: 
profile of the same scale; 13: inner view, detail of one cleared and stained specimen with 8 hooks, 


SL 56.0 mm, scale= 0.5 mm. 


254 MARIA DE LAS MERCEDES AZPELICUETA & JOSÉ O. GARCIA 


(Casciotta et al., 1992). Hooks are not common in the caudal fin, but, Fink & Weitzman 
(1974) described and illustrated hooks not only on the caudal fin of Saccoderma 
hastata but also on that fin of Cheirodon dialepturus. 

In glandulocaudine species, with notable sexual dimorphism, hooks may occur 
on caudal fin, as in Xenurobrycon macropus (Mahnert & Géry, 1984; Weitzman & 
Fink, 1985), X. pteropus, Scophaerocharax octopodus, Corynopoma risei, Gephyro- 
charax atricaudatus (Weitzman & Fink, 1985), Mimagoniates microlepis and M. 
reocharis (Menezes & Weitzman, 1990), Ptychocharax rhyacophila (Weitzman et al., 
1994) Tyttocharax cochui (Weitzman & Ortega, 1995). The presence of hooks in anal 
and pelvic fins is usual among glandulocuadine fishes; the position, size and shape of 
the hooks vary in different species. Mahnert & Géry (1984) mentioned the occurrence 
of hooks on the pectoral fin only in one specimen of X. macropus. 

Many other species of characins have been described with hooks; some of them 
have hooks on anal fins as Hyphessobrycon diancistrus (Weitzman, 1977a) and H. 
procerus (Mahnert & Géry, 1987) or hooks may be present on pelvic fins as in 
Paracheirodon axelrodi (Weitzman & Fink, 1983). In most of the species, hooks 
develop on anal and pelvic fins: Hyphessobrycon guarani (Mahnert & Géry, 1987), H. 
arianne (Uj & Géry, 1989), H. epicharis (Weitzman & Palmer, 1997), H. wajat 
(Almirén & Casciotta, 1999), Rachoviscus gracilipes and R. crassipes (Weitzman & 
Goncalves da Cruz, 1981), Piabarcus annalis and P. torrenticola, Creagrutus para- 
guayensis (Mahnert & Géry, 1988), some Trans-Andean species of Creagrutus as C. 
caucanus, C. maracaiboensis, C. paralacus or C. affinis (Harold & Vari, 1994), Hemi- 
grammus mahnerti (Uj & Géry, 1989), Bryconamericus theringi (pers. obs.). 

Well developed hooks occur also on anal or pelvic fins of other characiform 
species such as Brycon microlepis (Gery & Mahnert, 1992), the miniature Priocharax 
ariel (Weitzman & Vari, 1987), Toracocharax stellatus (pers. obs.) as well as in some 
species of different genera of the Stethaprioninae such as Poptella, Stethaprion, 
Brachychalcinus and Orthospinus (Reis, 1989). 

In the genus Jupiaba, J. meunieri or J. maroniensis (Gery et al., 1996, as 
Astyanax) show hooks on the anal fin. Within Astyanax, hooks on anal and pelvic fins 
have been reported in Astyanax maculisquamis (Garutti & Britski, 1997), A. alburnus 
(Malabarba, 1983 as A. hasemani), A. kullanderi (Costa, 1995), A. unitaeniatus 
(Garutti, 1998), A. eigenmanniorum, A. cf. fasciatus, A. cf. asuncionensis or A. alleni 
(pers. obs.). None of those species have caudal fin hooks. 

Extremely scarce is the information about the presence of hooks on dorsal fin; 
Weitzman (1977b) found hooks on that fin in two species of Hyphessobrycon only, the 
Amazonian H. socolofi and H. erythrostigma which bears very few hooks. 

The hooks of the pelvic axillary scale appear in medium sized males of A. ojiara 
and their number varies in different specimens, although one or two hooks are present 
usually (Figs 11, 12). One cleared and stained specimen has eight hooks (Fig. 13) on 
that scale. No other characids have been described with hooks on all fins or on pelvic 
axillary scale. 


A NEW SPECIES OF ASTYANAX FROM ARGENTINA 255 


TABLE | 


Morphometrics of holotype, 12 females and 12 males (paratypes) of Astyanax ojiara. Minimum, 
maximum and mean in parenthesis. DLAR-HJ= distance between last anal-fin ray and hypural 
joint. 


Holotype females males 

Standard length 50.5 39.8-72.0 38.0-53.5 
% of standard length 
Predorsal distance 51.4 51.8-56.9 (44.8) 50:9-595. 1183) 
Preventral distance 49.3 48.7-51.6 (50.4) 47.7-51.3 (48.8) 
Preanal distance 64.0 65.8-70.4 (67.7) 61.3-65.9 (63.4) 
Body depth 35.0 35.5-40.0 (38.1) 34.0-37.9 (36.5) 
Dorsal-fin base 13.4 13.8-17.2 (15.4) 12:8 = 15.9% (1323) 
Anal-fin base 32.4 27.6-33.4 (30.5) 30.0-34.6 (32.2) 
Pectoral-fin length 23.3 20.3-24.6 (21.7) 22.1-24.5 (23.2) 
Pelvic-fin length 19.6 14.8-18.3 (16.5) 17.9-21.5 (19.0) 
Distance between pectoral 

and pelvic fin origins 2255 20.4-25.7 (23.2) 21.8-24.6 (23.4) 
Distance between pelvic 

and anal fin origins 19:2 19.6-22.8 (21.3) 16.1-20.0 (18.2) 
Head length ZA, 27.3-29.5 (28.1) 26.5-29.7 (27.8) 
% of DLAR-HJ 
Peduncle depth 90.7 90.0-110.0 (99.0) 74.6-90.7 (83.0) 
% of head length 
DLAR-HJ 46.4 38.8-46.4 (42.3) 46.8-56.8 (50.2) 
Snout length 28.5 24.5-30.9 (27.7) 27.8-30.5 (29.2) 
Eye SIN 29.1-37.2 (28.1) 32.3-35.9 (34.3) 
Postorbital length SUA 50.0-54.8 (52.4) 53.4-57.2 (55.8) 
Interorbital length 32.8 31.0-34.6 (33.0) 31.0-35.6 (32.5) 
Maxillary length 22.8 21.1-24.5 (22.8) 37.1-40.6 (38.8) 
ACKNOWLEDGMENTS 


The authors thank L. R. Malabarba for exchange of specimens of A. eigen- 
maniorum, J. R. Casciotta for the photograph, A. Martinez Azpelicueta for the help 
with software, J. M. Diaz de Astarloa for making available some literature, and Consejo 
Nacional de Investigaciones Cientificas y Técnicas for financial support. 


REFERENCES 


ALMIRON, A. E. & CASCIOTTA, J. R. 1999. Hyphessobrycon wajat n. sp. from la Plata basin in 
Argentina (Characiformes: Characidae). Revue suisse de Zoologie 106: 339-346. 


AZPELICUETA, M. M. 1979. Anatomia comparada craneana y cintura pectoral de peces Chara- 
ciformes. Doctoral Thesis Nro. 372, Facultad de Ciencias Naturales y Museo, Univer- 
sidad Nacional de La Plata, 100 pp. 

BOHLKE, J. 1954. Studies on fishes of the family Characidae. N° 6. A synopsis of the Iguano- 
dectinae. Annals Magazine of Natural History, series 12, volume 7: 99-104. 

CASCIOTTA, J. R., MIQUELARENA, A. & PROTOGINO, L. 1992. A new species of Odontostilbe 
(Teleostei, Characidae) from the Uruguay Basin, with comments on the diagnostic 


characters of the genus. Occasional Papers, Museum of Natural History, Kansas 149: 
1-16. 


256 MARIA DE LAS MERCEDES AZPELICUETA & JOSÉ O. GARCIA 


Cope, E. D. 1876. On the fishes obtained by the Naturalist Expedition in Rio Grande do Sul. 
Proceedings of the American Transaction Philosophical Society 33: 84-89. 


Costa, W. J. E. M. 1995. Description of a new species of the genus Astyanax (Characiformes: 
Characidae) from the rio Araguaia basin, Brazil. Revue suisse de Zoologie 102: 257-262. 


EIGENMANN, C. H., 1921. The American Characidae. Memoirs of the Museum of Comparative 
Zoology, Harvard University 43: 209-310. 

EIGENMANN, C. H., 1927. The American Characidae. Memoirs of the Museum of Comparative 
Zoology, Harvard University: 311-428. 

FINK, W. L. & WEITZMAN, S. H. 1974. The so-called Cheirodontin fishes of Central America with 
descriptions of two new species (Pisces: Characidae). Smithsonian Contributions to 
Zoology 172: 1-46. 

GARUTTI, V. 1998. Descriçäo de uma espécie nova de Astyanax (Teleostei, Characidae) da bacia 
do rio Tocantins, Brasil. /heringia, serie Zoologia 85: 115-122. 


GARUTTI, V. & BRITSKI, H. A. 1997. Descriçäo de uma espécie nova de Astyanax (Teleostei, 
Characidae), com mancha umeral horizontalmente ovalada, da bacia do rio Guaporé, 
Amazonia. Papéis Avulsos de Zoologia, Sao Paulo 40: 217-229. 


GERY, J. & MAHNERT, V. 1992. Note sur quelques Brycon des bassins de I’ Amazone, du Parana- 
Paraguay et du Sud-Est brésilien (Pisces, Characiformes, Characidae). Revue suisse de 
Zoologie 99: 793-819. 


GERY, J., PLANQUETTE, P. & LE BAIL, P.-Y. 1996. Nouvelles espèces guyanaises d’Astyanax s.l. 
(Teleostei, Characiformes, Characidae) a épines pelviennes, avec une introduction 
concernant le groupe. Cybium 20: 3-36. 

HAROLD, A. S. & VARI, R. P. 1994. Systematics of the Trans-Andean species of Creagrutus 
(Ostariophysi: Characiformes: Characidae). Smithsonian Contributions to Zoology 551: 
1-31. 

Lucena, Z. M. & LUCENA, C. A. 1992. Revisäo das espécies do género Deuterodon Eigenmann, 
1907 dos sistemas costeiros do sul do Brasil com a descriçao de quatro espécies novas 
(Ostariophysi, Characiformes, Characidae). Comunicagöes do Museu de Ciéncias 
PUCRS, série zoologiia, Porto Alegre 5: 123-168. 

MALABARBA, L. R. 1983. Descriçäo e discussäo da posicäo taxonömica de Astyanax hasemani 
Eigenmann, 1914 (Teleostei, Characidae). Comunicacöes do Museu de Ciencias PUCRS, 
Porto Alegre 29: 177-199. 

MALABARBA, L. R. 1998. Monophyly of the Cheirodontinae, Characters and Major Clades 
(Ostariophysi, Characidae) (pp. 193-233). Jn: Malabarba, L. R., Reis, R. E., Vari, R. P., 
Lucena, Z. M., Lucena, C. A. (eds). Phylogeny and classification of Neotropical Fishes. 
Part 2. EdiPUCRS, Porto Alegre, 603 pp. 

MAHNERT, V. & GERY, J. 1984. Poissons Characoïdes (Characoidea) du Paraguay I: Xenuro- 
brycon macropus Myers et Miranda Ribeiro. Revue suisse de Zoologie 91: 497-513. 


MAHNERT, V. & GÉRY, J. 1987. Deux nouvelles espèces du genre Hyphessobrycon (Pisces, 
Ostariophysi, Characidae) du Paraguay: H. guarani Sp. n. et H. procerus Sp. n. Bonner 
zoologische Beiträge 38: 307-314. 

MAHNERT, V. & GÉRY, J. 1988. Les genres Piabarchus Myers et Creagrutus Günther du 
Paraguay, avec la description de deux nouvelles espèces (Pisces, Ostariophysi, Chara- 
cidae). Revue française d’Aquariologie 15: 1-8. 

Menezes, N. A. & WEITZMAN, S. H. 1990. Two new species of Mimagoniates (Teleostei: 
Characidae: Glandulocaudinae), their phylogeny and biogeography and a key to the glan- 
dulocaudin fishes of Brazil and Paraguay. Proceedings of the Biological Society of 
Washington 103: 380-426. 

Reis, R. E. 1989. Systematic revision of the Neotropical characid subfamily Stethaprioninae 
(Pisces, Characiformes). Comunicacöes do Museu de Ciéncias PUCRS, serie zoologia, 
Porto Alegre 2: 3-86. 


A NEW SPECIES OF ASTYANAX FROM ARGENTINA 257 


TAYLOR, W. R. & VAN Dyke, G. C. 1985. Revised procedures for staining and clearing small 
fishes and other vertebrates for bone and cartilage study. Cybium 9: 107-119. 

Us, A. 1987. Les Cheirodontinae (Characidae, Ostariophysi) du Paraguay. Revue suisse de 
Zoologie 94: 129-175. 

Ur, A. & Gery, J. 1989. Deux nouvelles espèces de Tetras (Poissons characoides, Characidae 
auct., Tetragonopterinae) du Paraguay: Hyphessobrycon arianne sp. n. et Hemigrammus 
mahnerti sp. n. Revue suisse de Zoologie 96: 147-159. 

VARI, R. P. & GERY, J. 1980. Cheirodon ortegai, a new markedly sexually dimorphic 
cheirodontine (Pisces: Characoidea) from the rio Ucayali of Peri. Proceedings of the 
Biological Society of Washington 93: 75-82. 

WEITZMAN, S. H. 1977a. A new species of characoid fish, Hyphessobrycon diancistrus, from the 
rio Vichada, Orinoco river drainage, Colombia, South America (Teleostei: Characidae). 
Proceedings of the Biological Society of Washington 90: 348-357. 


WEITZMAN, S. H. 1977b. Hyphessobrycon socolofi, a new species of characoid fish (Teleostei: 
Characidae) from the rio Negro of Brazil. Proceedings of the Biological Society of 
Washington 90: 326-347. 

WEITZMAN, S. H. & THOMERSON, J. E. 1970. A new species of glandulocaudine characid fish, 
Hysteronotus myersi, from Pert. Proceedings of the California Academy of Sciences 38: 


139-156. 


WEITZMAN, S. H. & GONCALVES DA CRUZ, C. A. 1981. The South American fish genus 
Rachoviscus, with a description of a new species (Teleostei, Characidae). Proceedings of 
the Biological Society of Washington 93: 997-1015. 


WEITZMAN, S. H. & FINK, W. L. 1983. Relationships of the Neon Tetras, a group of South 
American freshwater fishes (Teleostei, Characidae) with comments on the phylogeny of 
the New World Characiforms. Bulletin of the Museum of Comparative Zoology, Harvard 
150: 339-395. 


WEITZMAN, S. H. & FINK, S. V. 1985. Xenurobryconin phylogeny and putative pheromone pumps 
in Glandulocaudine fishes (Teleostei: Characidae). Smithsonian Contributions to Zoology 
421: 1-121. 

WEITZMAN, S. H. & VARI, R. P. 1987. Two new species and new genus of miniature characid 
fishes (Teleostei: Characiformes) from Northern South America. Proceedings of the 
Biological Society of Washington 100: 640-652. 

WEITZMAN, S. H. & ORTEGA, H. 1995. A new species of Tyttocharax (Teleostei: Characidae: 
Glandulocaudinae: Xenurobryconini) from the rio Madre de Dios basin of Perù. /chthyo- 
logical Exploration of Freshwaters 6: 129-148. 


WEITZMAN, S. H. & PALMER, L. 1997. A new species of Hyphessobrycon (Teleostei: Characidae) 
from the Neblina region of Venezuela and Brazil, with comments on the putative ‘rosy 
tetra clade’. Ichthyological Exploration of Freshwaters 7: 209-242. 

WEITZMAN, S. H., FINK, S. V., MACHADO-ALLISON, A. & ROYERO L., R. 1994. A new genus and 
species of Glandulocaudine (Teleostei: Characidae) from Southern Venezuela. /chthyo- 
logical Exploration of Freshwaters 5: 45-64. 

ZANATA, A. M. 1997. Jupiaba, um novo género de Tetragonopterinae com osso pélvico em forma 


de espinho (Characidae, Characiformes). /heringia, Serie Zoologia, Porto Alegre 83: 
99-136. 


' N 
FR hy 


ey 


my A MSHS 


| À i f 17 
vie a iron | 
fit ibi “ FR PEUT E L 
cy eal re be | 


12 


REVUE SUISSE DE ZOOLOGIE 107 (2): 259-275; juin 2000 


Revision of the genus Ebbrittoniella Martinez 
(Coleoptera: Scarabaeoidea: Ceratocanthidae)! 


Alberto BALLERIO 
Viale Venezia 45, I-25123 Brescia, Italy. 
E-mail: ballerio@numerica.it 


Revision of the genus Ebbrittoniella Martinez (Coleoptera: Scarabae- 
oidea: Ceratocanthidae). - The Oriental genus Ebbrittoniella Martinez, 
1962 is redescribed. The current spelling Eubrittoniella is emended. 
Cyphopisthes gestroi (Paulian, 1942) is transferred to Ebbrittoniella. 
Presently the genus includes E. ignita (Westwood, 1883) and E. gestroi 
(Paulian, 1942). Some features of male and female genitalia and the 
affinities of the genus are briefly discussed. 


Key-words: Coleoptera - Scarabaeoidea - Ceratocanthidae - Ebbrittoniella 
- Taxonomy - Oriental Region. 


INTRODUCTION 


In 1883 Westwood described Acanthocerus (Sphaeromorphus) ignitus on the 
basis of a specimen from Sumatra. In the subsequent fifty years Preudhomme de Borre 
(1886), Lansberge (1887), Gestro (1899) and Arrow (1912) dealt with this species; 
among these authors Gestro for the first time stressed the unnatural generic placement 
of the taxon, being the only Oriental representative in a New World genus. In 1962 
Martinez, by examining a specimen from Sarawak in BMNH, followed Gestro’s 
suggestion and created the new genus Ebbrittoniella to accommodate the taxon ignitus, 
while pointing out that the closest genus to it was the Neotropical genus Acanthocerus 
Macleay, 1819 (now Ceratocanthus White, 1842). 

Paulian in his revision of Oriental Ceratocanthidae (1978) summarized available 
distributional data, redescribed the species and placed it in the key near Cyphopisthes 
Gestro, 1899. He also emended the name in Eubrittoniella ignitus. This emendation 
cannot be considered justified according to the art. 33 of ICZN, because in the original 
paper there is no clear evidence of an author’s lapsus calami or printer’s error as 
defined by art. 32 of ICZN and therefore the original spelling Ebbrittoniella should be 
maintained. 

After that point the genus Ebbrittoniella has been listed only by Browne & 
Scholtz (1995, 1996, 1998) as Eubrittoniella, in a series of papers upon the hindwing 
articulation, base and venation of the Scarabaeoidea. 


! Sth contribution to the knowledge of Ceratocanthidae. 
Manuscript accepted 28.02.2000 


260 ALBERTO BALLERIO 


During revisionary studies on Old World Ceratocanthidae I had the opportunity 
to examine the type material as well as specimens from several collections, allowing 
me to confirm the validity of the genus. Cyphopisthes gestroi (Paulian, 1942) shares all 
the generic characters and therefore it is now transferred to Ebbrittoniella. 


METHODS AND ACRONYMS 


Elytral length is measured from the apex of elytral articular process to the more 
external point of the apical convexity, total width measurements are the sum of both 
elytral widths. 

Drawings of genitalia were taken from pieces previously cleaned with 10% 
KOH solution. 

Terminology follows Nel & De Villiers (1988) and Nel & Scholtz (1990) for 
mouthparts, Kukalovä-Peck & Lawrence (1993) for wing venation and D’Hotman & 
Scholtz (1990) for male genitalia (thus the dorsal side of aedeagus is the concave one); 
for the remaining conventions I refer to Ballerio (2000). 


EE maximum elytral length 
EW maximum total elytral width 
HL maximum head length 

HW maximum head width 

È length 

PL maximum pronotum length 
PW maximum pronotum width 
W width 


ABCB A. Ballerio private collection, Brescia (Italy) 

BMNH The Natural History Museum, London 

BPBM Bernice Bishop Museum, Honolulu 

HNHM Hungarian Natural History Museum, Budapest 

MCSN Museo Civico di Storia Naturale “Giacomo Doria”, Genova 
MHNG Muséum d’histoire naturelle, Genève 

MNHN Musée national d'Histoire naturelle, Paris 

NHMW Naturhistorisches Museum, Wien 

NMPC National Museum (Natural History), Praha 

OXUM Hope Department of Entomology, Oxford University, Oxford 
RMNH Nationaal Natuurhistorisch Museum, Leiden 

SACF _ S. Adebratt private collection, Frinnaryd/Boxholm (Sweden) 
SMTD Staatliches Museum für Tierkunde, Dresden 

ZMUC Zoologisk Museum, Kobenhavns Universitet, Kobenhavn 


TAXONOMY 


Ebbrittoniella Martinez 


Ebbrittoniella Martinez, 1962: 61 (description) 
Eubrittoniella Martinez, Paulian, 1978: 506 (emendation) 


REVISION OF THE GENUS EBBRITTONIELLA 261 


Type species: Acanthocerus (Sphaeromorphus) ignitus Westwood, 1883 by ori- 
ginal monotypy. 

Etymology: named after Dr. E. B. Britton, former curator at BMNH. The gender 
is feminine. 


DIAGNOSIS 


The genus can be identified by the following combination of characters: labrum 
distally abruptly truncate, truncature marked dorsally by a slight carina bearing a row of . 
long, erect, fine setae, truncature in frontal view forming a plate irregularly semi- 
circular, with semicircumference corresponding to the carina; genal canthus complete, 
touching the occipital area; dorsal ocular area large; anterior angles of pronotum 
broadly rounded; mesotibiae short and wide (W/L ratio = 0.3), in dd with the inner 
apical spur straight and very short; protibiae with outer edge almost smooth (low 
magnification), ending with a single tooth in both sexes; apical spur of protibiae sharp 
and slender. 


DESCRIPTION 


Medium to large Ceratocanthidae; volant. 

Head: W/L ratio = 1.8, subpentagonal, anterior edge forming a triangle with 
obtuse apex (about 150°), both sides of the triangle smooth and almost rectilinear, not 
reflexed upward; genae aligned with fore edge, forming a right angle with genal 
canthus; genal canthus narrow, straight and complete, touching the occipital area; dor- 
sal ocular area large; dorsal interocular area about five times the maximum width of the 
dorsal ocular area; ventral ocular area very large; head surface almost plane. 

Pronotum:W/L ratio = 1.8, slightly wider than maximum elytral width, evenly 
and broadly convex; anterior edge feebly bisinuate; anterior angles distinctly but 
slightly protrudent forward, broadly and regularly rounded; sides obtusely rounded; 
base narrow, a very thin bead present anteriorly and at each side of base; base with a 
weak callosity marking each extremity of base of scutellum. 

Scutellum: very large, longer than wide (W/L ratio = 0.7), sides proximally 
subparallel and distinctly notched by apical portion of mesepisternum and elytral 
articular process, then convergent to form a triangle with apex very elongate and acute 
and sides slightly curved inward; apical portion of mesepisterna (see Ballerio, 2000) 
visible from above, very large, larger than elytral articular process, subrectangular, 
smooth and shiny (Fig. 2d). 

Elytra: slightly longer than wide (W/L ratio = 0.9), maximum width near 
middle, apex in lateral view fairly reentering inward; slightly flattened on disc, then 
abruptly convex at sides to form a pseudoepipleuron not marked by any lateral carina; 
marginal elytral area narrow, almost indistinct; elytral suture very feebly raised; sutural 
stria very fine and close to suture, limited to distal third; inferior sutural stria absent; 
striated articular area well developed and visible in lateral view, relatively wide and 
long; elytral articular process well developed, smooth and shiny. 

Apical extremity of clypeus (see Ballerio, 2000) short and transversely grooved. 
Labrum (Fig. 4b) wide and short, proximally with surface microreticulated, distally 


262 ALBERTO BALLERIO 


abruptly truncate, truncature marked dorsally by a slight carina bearing a row of long, 
erect, distally curled, fine setae; truncature in frontal view forming a plate irregularly 
semicircular, with semicircumference corresponding to the carina; surface of plate 
almost smooth, bearing few long setae. Distal epipharynx longitudinally divided by a 
very sharp strong anterior median process, distally very raised; median brush and 
corypha absent; apical fringe made of long fine setae, absent in the middle; lateral 
combs made of long fine setae. Mentum ventrally flat, deeply emarginated in the 
middle of anterior edge, emargination regularly wide-U-shaped; labial palpi (including 
palpiger) four jointed, first joint short and transverse, joint two securiform, joint three 
short and ovoidal, the same length of the second, and joint four subconical, about two 
times the length of the former, all joints, apart from the last one, fringed with long 
setae. Maxillae (Fig. 3a) with a very elongate single lacinia, covered with fine long 
setae, monolobed galea proximally sclerotized and distally clothed with very coarse 
short thick bi- or triphid setae (galeal brush) (Fig. 3b); maxillary palpi (including 
palpiger) four jointed, palpiger very small, joint two wide and relatively short, distinctly 
wider than the following joints, joint three about as wide as long, joint four long and 
subconical, slightly longer than the preceding two together, apically bearing some short 
sensilla. Mandibles elongate, slightly asymmetrical, apicalis more or less gently bent at 
about a right angle with apex short and acutely pointed, mesal brush narrow and well 
developed, conjunctive present, molar lobe very strong. Antennae 10-jointed, scape 
large, distally subcarinate (securiform), distally bearing some setae, funicle short with 
pedicellum plump and rounded, the remaining joints very short, distinctly wider than 
long, antennal club three-jointed, joints hairy, relatively short, narrow; club small, about 
as long as wide and as wide as the length of funicle (L funicle/L lamellae ratio = 2.1). 
Ventral areas of prothorax: (Fig. 4a) sides of propleura smooth, very excavated 
and folded in, excavation with a further shallow narrow reniform excavation inside 
(visible also in dorsal view against the light as a dark reniform patch). Procoxae 
transversely oriented, apices nearly touching each other. Anterior trochanters with 
anterior tips bearing a tuft of long setae. Profemora slender (W/L ratio = 0.2), posterior 
edge without emargination, surface smooth. Protibiae almost straight, outer edge 
smooth at low magnfication (at most very slightly serrate: few feeble denticles visible 
at 45x); apical spur relatively short, sharp, very gently and feebly curved downward. 
Protarsi with first article as long as the following three together, articles two and three 
slightly dilated, article five slightly longer than the former; each tarsomere, with the 
exception of the last one, ventrally bearing a tuft of dense fine setae. Mesosternum 
forming a sharp fine carina, protruding between mesocoxae and joining metasternum. 
Mesocoxae almost adjacent to each other, longitudinally oriented. Trochanters with 
acute posterior tip. Mesofemora slender (W/L ratio = 0.3), surface smooth, with 
posterior edge emarginated at distal third, emargination preceded by a small distinct 
tooth. Mesotibiae subrectangular, short (W/L ratio = 0.3), inner angle of apex with two 
straight apical spurs. Mesotarsi inserted near the inner angle of apical edge, slightly 
longer than apical edge of tibia (exceding it for the length of the last tarsomere), with 
first three articles subequal in length, fourth shorter, fifth almost as long as the 
preceding two; each tarsomere, with the exception of the last one, ventrally bearing a 


263 


REVISION OF THE GENUS EBBRITTONIELLA 


UOIQUIE I) JINPe JO snyrgey 


€ 


(ueipneg) 10.415938 D]]21uon11gQqg :q ‘(WiOref eyeT) impe 


(ur 
Jo smiqey 


€ 


Jeq a[eos) (APIAGOM 


f Aq ssurmesp) (spuejy3iH 


(POOMISOM) DHUS1 DJJ21U01149q4 8 — | ‘OM 


264 ALBERTO BALLERIO 


tuft of dense setae. Trochanters of metafemora with acute posterior tip, metafemora 
plumper than mesofemora (W/L ratio = 0.3), surface wrinkled, posterior edge distally 
with a small emargination. Metatibiae triangular, wide (W/L ratio = 0.5), ending with 
two straight sharp fine paired spurs. Metatarsi almost as long as the apical edge of tibia, 
first article almost as long as the following three together, fifth almost as long as the 
first one; each tarsomere, with the exception of the last one, ventrally bearing a tuft of 
dense setae. 

Wings (Fig. 2a) (Lwing/Lelytron ratio = 2.1): fully developed, MP, ,, - RP loop 
present with RP long (although weakly sclerotized), MP, medium sized, apical field 
with a vertical secondary sclerification near the radial cell. First axillary with no 
appreciable differences at species level (Fig. 2b). 

Sexual dimorphism: 2 9 (Fig. 2c: B) have the apical outer tooth of protibiae 
distinctly sharper and more protruding outward and forward than in dg d (Fig. 2c: A) 
and mesotibiae with the inner apical spur straight and approximately as long as the 
outer, while in d d it is straight but very short and very difficult to detect. 

Male genitalia: Genital segment (Fig. 5c) Y-shaped (no appreciable differences 
at species level), fairly sclerotized, with a distinct manubrium about as long as the basal 
triangle, branches forming the manubrium apparently not fused together, although 
connected by a transparent membrane (after treatment with KOH), base of triangle very 
weakly sclerotized; basal piece of aedeagus large and twisted (Fig. 5a), about four times 
the length of parameres; internal sac very large about three times as long as tegmen, 
internal sac distally with coarse spicules and setae; temones present; parameres (Fig. 
5b) short and slightly asymmetrical (hardly appreciable differences at species level), 
laterally flattened, between parameres dorsally lies a narrow subtriangular sclerite. 

Female genitalia: bursa copulatrix (Fig. 5d) with one small symmetrical sub- 
circular or subtriangular sclerite (very variable in shape and without appreciable 
differences at species level) with a hole in the middle; spermatheca (Fig. 6) strongly 
sclerotized, large and distinctly wide-U-shaped; genital palpi weakly sclerotized, sub- 
circular, relatively small and short. 


Ebbrittoniella ignita (Westwood, 1883) Fig. la 


Acanthocerus (Sphaeromorphus) ignitus Westwood, 1883: 2 (description) 

Acanthocerus (Sphaeromorphus) ignitus Westwood: Preudhomme de Borre, 1886: 80 (catalogue) 

Synarmostes ignitus (Westwood): Lansberge, 1887: 209 (list) 

Acanthocerus ignitus Westwood: Gestro, 1899: 462 (redescription, distribution and key) 

Acanthocerus ignitus Westwood: Arrow, 1912: 49 (catalogue) 

Ebbrittoniella ignita (Westwood): Martinez, 1962: 61 (redescription and iconography) 

Eubrittoniella ignitus (Westwood): Paulian, 1978: 506 (redescription, distribution, iconography 
and key) 

Type locality: Koetoer (Indonesia: Sumatra). 

Material examined - Holotypus d: Koetoer 6.78 / Sum. Exp. Koetoer 6.78 / Typus / 
Acanthocerus ignitus Westw. / Museum Leiden Acanthocerus ignitus det. Westwood / 6903 / 
Eubrittoniella ignitus (Westw.) R. Paulian det. / Ebbrittoniella ignita (Westw.) A. Ballerio det. 
1997 (RMNH), [very damaged specimen, pinned, lacking the head, which is glued together with 
legs and mouthparts on a card pinned with a separate pin, without any data bearing label]. 


REVISION OF THE GENUS EBBRITTONIELLA 265 


b Coi 


A 


Fic. 2 — a: E. gestroi (Cameron Highlands), wing: vertical secondary sclerification (S) (scale bar: 
1 mm); b: E. gestroi (Cameron Highlands), first axillary (scale bar: 0,1 mm; c: Ebbrittoniella, 
apex of foretibiae (dorsal view): (A) male, (B) female; d: Ebbrittoniella, area near scutellum as 
seen when the beetle assumes the “rolled up” posture: apical portion of mesepistemum (M), 
articular process of elytron (AP), scutellum (S), elytron (E), pronotum (P). 


Other 32 specimens (7 dd and 4 9 2 dissected). INDONESIA. Kajoe Tanam [Sumatra], 
Klein (ZMUC); Sumatra, Manna, M. Knapper (RMNH); Ruuyer Payakor, Sumatra (RMNH); 
Sentinjak, Sumatra, 1800 ft., Jan. ‘98, 99-95 (BMNH); Setinjak, W. Sumatra, I to VII. 98, 
(Ericsson) (Coll. C. Felsche, Kauf 20, 1918) (SMTD); Sumatra, Padang Sidempoean, XII.1902- 
1.1903 (MNHM); Sumatra (ex Mus. Van Lansberge) (MNHM). Nias Island. Is. Nias, U. Raap 
(MNHM); Hili Madjedja, N. Nias, 4me trim. 195, I.Z. Kannegieter (ex Mus. Van de Poll) 
(MNHM); Is. Nias 1897-98, U. Raap (RMNH); Is. Nias 1897-98 U. Raap (MCSN). Kalimantan- 
Timur: Apokayan, Long Sungei Barang 900 m, secondary forest, 15-23.02.1997, leg. C. & P. 
Zorn. MALAYSIA. Malay penin.: Selangor, Giunting Simpak, Jan. 2"¢ 1933, N.M. Pendlebury, 


266 ALBERTO BALLERIO 


F.M.S. Museum (BMNH); Malaysia: Pahang/Johor, Endau Rompin NP, 100 m, Salendang, 28.2- 
12.3 1995, leg. Strba & Hergovits (NHMW); Malaysia — Pahang, Banjaran Benom, Lata Jarom, 
6-8.3.1997, Ivo Jenis leg. (ABCB); ibidem, 18-21.3.1997, Ivo Jenis leg. (ABCB), W. Malaysia, 
Pahang, Baniaran Benom Mts., K. Ulu Dong 10-15 km SSE, 17-23.1V.1997, D. Hauck leg. 
(ABCB); Sarawak, C. S. Brooks, B.M. 1928-193 (BMNH); Malaysia: Sabah, Sipitang, Mendo- 
long, T1B/W4, 11.V.1988, leg. S. Adebratt, 12-60, 2859BC (SACF). OTHER. Sunda-Inseln v. 
Studt /coll. Petrovitz (MHNG). 

DESCRIPTION 

HL= 0.8-1.2 mm HW= 1.6-2 mm PL= 1.5-2 mm PW=2.8-3.3 mm EL= 2.9-3.9 mm 
EW= 2.7-3.6 mm. 

Dorsally very shiny and entirely brightly metallic: red with gold/green faint, 
elytral suture, scutellum, sides of elytra, sides of pronotum and sides of head deep 
electric blue with green faint; ventrally alutaceous, reddish brown, antennae yellowish. 
Pubescence invisible at low magnification, otherwise formed by very short and fine 
hyaline erect setae, subject to wear. Head surface with a variable number of transverse 
lines, occupying anteriorly the two thirds of head; vertex with sparse simple punctures, 
often preceded by a very short transversal impressed line. Pronotum very convex, 
surface with very small horseshoe-shaped punctures (containing a small simple 
puncture in the middle) or sometimes simple punctures, never very dense, usually 
sparser on disc and slightly coarser near anterior angles; the horseshoe is opened 
outwards in the punctures at sides and inwards in the punctures on the disc. Scutellum 
with horseshoe-shaped punctures with horseshoe opened toward apex, punctures denser 
near sides. Elytra distinctly convex, surface with punctures horseshoe-shaped, small, 
never very dense, variably distributed, horseshoe opened toward suture or apex; two 
longitudinal lines along the suture at medial and distal third of length; pseudoepipleura 
with some longitudinal lines; humeral callus not protruding; articular area very large. 
Outer face of meso- and metatibiae with short impressed lines varying from longitu- 
dinal to transverse and short erect setae through the entire surface. Wings with vertical 
secondary sclerification of apical field finer than in E. gestroi and MP,,, - RP loop 
shorter. Spermatheca as in figs. 6a, b, c, d. 


VARIABILITY 


Although the majority of individuals examined shows the typical colours, one 
pair from Pahang is uniformly dark green, one individual from Nias is uniformly deep 
amaranth and the individual from Sabah is yellowish with greenish metallic sheen; 
there is a strong variability also in the punctures (mostly the ones of pronotum), which 
vary in size and density: the specimen from Sabah has the entire head surface covered 
by transverse lines, the amaranth specimen from Nias has the entire pronotum 
completely smooth, while the pair of dark green specimens has punctures coarser and 
more impressed than the other specimens. The very fine pubescence varies and in 
some specimens is no longer visible (probably ripped off by wear). 


DISTRIBUTION AND HABITAT 


Recorded from Peninsular Malaysia, Sumatra, Nias Island and Borneo. For 
habitat see under E. gestroi. 


REVISION OF THE GENUS EBBRITTONIELLA 267 


Fic. 3 — a: E. gestroi (Cameron Highlands), maxilla (SEM micrograph); b: E. gestroi (Cameron 
Highlands), detail of galeal brush (SEM micrograph). 


268 ALBERTO BALLERIO 


REMARKS 


°° 


Westwood (1883) described also a variety “paullo minor, obscurior,...” from 
Sarawak without naming it; examination of the specimen in OXUM revealed that it is 
an Eusphaeropeltis sp. 


Ebbrittoniella gestroi (Paulian, 1942) comb. n. Fig. 1b 


Philharmostes Gestroi Paulian, 1942: 70 (description and key) 
Cyphopisthes gestroi (Paulian): Paulian, 1978: 512 (new combination, redescription, distribution, 
iconography and key) 

Type locality: Palembang (Indonesia: Sumatra). 

Etymology: named after Dr. R. Gestro, late director of MCSN and specialist of 
Ceratocanthidae. 

Material examined — Holotypus, 2: Sumatra Palembang / Type / Cyphopisthes gestroi n. 
sp. det. R. Paulian, 1937/ Ebbrittoniella gestroi (Paulian) n. comb. det. A. Ballerio, 1997. 
(MNHN) [completely rolled up and glued on a card] . 

Other 141 specimens ( 15 dd and 15 9 9 dissected) - INDONESIA. Palembang 
[Sumatra], 1900, Bouchard (MCSN); MALAYSIA. Malaya: (W) Perak, Maxwell Hill, 1350 m., 
17-20.1IL.1958, T.C.Maa collector, Bishop (BPBM); Malaysia — Perak, Banjaran Bintang, 
Maxwell Hill (Taiping), 18-19.2.1997, leg. Ivo Jenis (ABCB); West Malaysia, Perak, Maxwell 
Hill, 900-1000 m, above Taiping City, 12-16.1.1995, leg. S. Beévar j. & s. (ABCB); W. 
Malaysia, Perak: 25 km NE of Ipoh, 2100 m, Banjaran Titi Wangsa mts., Gunung Korbu, 4- 
13.111.1998, P. Cechovskÿ leg.; W. Malaysia, Perak, 25 km NE of Ipoh, 1200 m, Banjaran Titi 
Wangsa mts., Gunung Korbu, 27.1-2.11.1999, P. Cechovsky leg.; W. Malaysia, Pahang, 30 km E 
of Ipoh, 1500 m, Cameron Highlands, Tanah Rata, 20.11-3.1II. 1998, P. Cechovsky leg.; ibidem, 
22-26.1.1999, P. Cechovsky leg.; P. Malaysia-Pahang: Cameron Highlands, Tanah Rata umg., 
gn. Jasar, 1300 m, 25.11.1997, leg. Schuh & Lang (ABCB); West Malaysia, Pahang, C. 
Highlands, Tanah rata, 20-25.1.1995, Gn. Jasar, 14-1500 m, leg. S. Beëvar j. & s. (ABCB); West 
Malaysia, Pahang, Cameron Highlands, Brinchang, 18-19.1.1995, Gunung Berembam, 1600 m, 
leg. S. Beévar j. & s. (ABCB); Malaysia, Pahang, Cameron Highlands, 2 km S of Tanah Rata on 
Tapah Road, montane rainforest, at light no. 93, 29.11.1995, leg. O. Merkl & L. Szikossy 
(HNHM); Malaysia, Pahang, Cameron Highlands, Tanah Rata, from illuminated white washed 
walls no. 77, 23-31.111.1995, 1. O. Merkl (HNHM); Malaysia — Pahang, Banjaram Benom, Lata 
Jarom, 6-8.3.1997, leg. Ivo Jeni$ (ABCB); Malaysia — Perak, Cameron Highlands, Tanah Rata, 
13-16.3.1997, leg. Ivo Jenis (ABCB); Borneo (Brit. N.), Sandakan bay (SW), Sapagaya Lumber 
Camp, 2-20 m, XI-3-157, J.L. Gressit Collector (BPBM); North Borneo (SE), Forest Camp, .19 
km N. of Kalabakan, 18.X1.1962, Y. Hirashima, Light trap, Bishop (BPBM); ibidem, 7- 
10.X1.1962 (BPBM); ibidem, 27.X.1962 (BPBM): ibidem, 10.X.1962 (BPBM); Malaysia — 
Sabah, Crocker Range National park, Longkogungan env., Ca. 750-850 m a.s.l., 19-21.VI.1996, 
7c (NHMW); Borneo: Sarawak Bau District, Bidi, 90-240 m. 3.IX.1958, T.C. Maa Collector 
Bishop (BPBM); Sarawak, Kapit dist., Sebong, Baleh riv., 9-21.3.1994, Sv. Bily leg. (NMPC), 
Sarawak, Kapit dist., Rumah Ugap vill., Sut. Riv., 3-9.3.1994, Sv. Bily leg. (NMPC); Sarawak: 
Gunung Mulu Nat. Park, R.G.S. Exped. 1977-8, J.D. Holloway et al. B.M. 1978-206, site 2, 
january, camp 4, Mulu, 1790 m, 452463, lower montane (moss) forest, acl-understorey (BMNH). 


DESCRIPTION: 
HL= 0.9-1.1 mm HW= 1.5-1.9 mm PL= 1.3-1.8 mm PW=2.7-3.2 mm EL= 3-3.5 mm 
EME 232 mms: 

Dorsally metallic bronze/green with reddish faint, head, pronotum, scutellum 
and elytra covered by long recumbent yellowish setae; ventrally alutaceous yellowish/ 
brown with antennae yellowish. Head as in E. ignita but completely covered by 
recumbent setae and dense punctures. Pronotum slightly less convex than in E. ignita, 


REVISION OF THE GENUS EBBRITTONIELLA 269 


FIG. 4 — a: E. gestroi (Cameron Highlands): ventral areas of prothorax (SEM micrograph); b: E. 
gestroi (Cameron Highlands): head in fronto-lateral view, showing the truncature of labrum 
(SEM micrograph). 


270 ALBERTO BALLERIO 


surface completely covered by very dense small horseshoe-shaped punctures, with 
horseshoe very short and almost transversal; each horseshoe with a small impressed 
puncture in the middle; pubescence very long and dense, recumbent, some subcircular 
small and weak depressions at each side of disc. Scutellum with coarse horseshoe- 
shaped punctures and long recumbent pubescence. Elytra slightly less convex than in 
E. ignita, surface with punctures less dense and slightly larger than on pronotum, 
relatively sparser near suture; proximal third with large longitudinal smooth areas 
between the suture and the pseudoepipleura; pseudoepipleura with transversal long 
dense horseshoe-shaped punctures. Humeral callus very developed and protuding 
outwards; articular area smaller than in FE. ignita. Outer face of meso- and metatibiae 
with short impressed lines varying from longitudinal to transverse and short erect setae 
through the entire surface. Wings (Fig. 2a): vertical secondary sclerification of apical 
field thicker than in E. ignita and MP, ,, - RP loop distinctly longer. Spermatheca as in 
figs. 6e, f, g, h. 


VARIABILITY 


E. gestroi is less variable compared to the former species; the colours vary from 
bronze green to green with faint reddish sheen. In the pair from Lata Jarom the smooth 
elytral areas are very small, while punctures and lines are slightly more impressed. 


DISTRIBUTION AND HABITAT 


Recorded from Peninsular Malaysia, Sumatra and Borneo (Sabah and Sarawak). 

Both species are found in rainforests, they are sympatric and sometimes also 
syntopic (Lata Jarom, leg. Jeni$). E. ignita seems to be restricted to lowland rainforests, 
where it has been collected by beating leaves or with window traps; E. gestroi shows a 
broader ecological range because it is found in both lowland and montane forests (till 
2100 m a.s.l.), where it is the most commonly collected Ceratocanthidae. There are 
several records of this species at light. The unusual vestiture of galeal brush, which is 
covered by short and thick bi- or triphids dense setae (while in most other Cerato- 
canthidae these setae are longer, finer and usually sharp) could suggest that Ebbritto- 
niella has feeding habits different from the ones of the majority of Ceratocanthidae. 


KEY TO THE SPECIES OF EBBRITTONIELLA 


I Labrum distally distinctly truncate, truncature marked dorsally by a 

slight carina bearing a row of long, fine, erect setae, truncature in frontal 

view forming a plate irregularly elliptical or semicircular................. 2 
- Labrum without distinct truncature marked dorsally by a carina 

DD RE ease uae 20 UNS NEE RISE ERE NEN INA i other Ceratocanthidae 
2 Anterior angles of pronotum triangular; mesotibiae slender and relatively 

narrow (W/L ratio = 0,2), with inner apical spur of d d bent inwards at a 

right angle; apex of protibiae usually ending with two teeth in the 9 9 

(one known exception); apical spur of protibiae strong, apically distinctly 

Pent downwards ohio) Met ee ee ee eee Cyphopisthes Gestro 


REVISION OF THE GENUS EBBRITTONIELLA 271 


- Anterior angles of pronotum broadly rounded; mesotibiae short and wide 
(W/L ratio = 0,3), with the inner apical spur of d d straight and extre- 
mely short; apex of protibiae ending with a single tooth in both sexes; 
apical spur of protibiae fine, very gently and regularly bent downwards. 
TT SE Ce me Sy tae ao Ebbrittoniella Martinez 
3 Surface of pronotum strongly punctured, punctures very dense, giving a 
granulose appearance, pronotal pubescence long, yellowish, recumbent, 
elytra with the same microsculpture and pubescence as pronotum, but 
with some smooth longitudinal areas, humeral callus very pronounced 
WET CRIE E. gestroi (Paulian) 
- Surface of pronotum smooth, with sparse simple or horseshoe shaped 
punctures, glabrous or at most with very fine and short erect hyaline 
pubescence, elytra like pronotum, humeral callus weakly pronounced 
De D MR SR RS I AE NER: E. ignita (Westwood) 


DISCUSSION 


Some interesting new discoveries about the genitalia of the Ceratocanthidae 
were made during this revision. First of all it was found that the bursa copulatrix, which 
contains the spermatophore, bears a small subcircular sclerite on its inner wall, which 
probably has the function of breaking the spermatophore; similar sclerites have been 
observed in other genera of Ceratocanthidae, namely Pterorthochaetes Gestro, 1899 
(Ballerio, 1999), Philharmostes Kolbe, 1895 and its allies (Ballerio, 2000) and 
Eusphaeropeltis Gestro, 1899 (author’s unpublished data). Secondly a mobile acces- 
sory sclerite was detected, lying between the ventral basal extremities of parameres and 
associated with the median lobe; normally it lies parallel to the anchoring point of 
parameres with basal piece, but, when the internal sac is everted, the sclerite is raised at 
right angle, perpendicular to the anchoring point of parameres; it is possibly an 
intermediate form between an anchoring and a supporting sclerite (see D’Hotman & 
Scholtz, 1990). 

In order to find the closest relationships of the genus Ebbrittoniella an analysis 
was conducted on all the genera of typical Ceratocanthidae, on the basis of all available 
morphological characters of adults, using the sister group of Ceratocanthidae, i.e. the 
Hybosoridae (Browne & Scholtz, 1996) and in particular the genera Phaechrous 
Castelnau, 1840 and Liparochrus Erichson, 1848 (Howden & Gill, 1995), as an out 
group for character polarization. The analysis suggests that Cyphopisthes Gestro, 1899 
could be the genus closest to Ebbrittoniella. The following synapomorphies are shared 
by the two genera: a) outer edge of protibiae smooth (low magnification), b) sexual 
dimorphism involving the apical teeth of protibiae, c) labrum distally truncate, trun- 
cature marked dorsally by a slight carina bearing a row of erect fine setae, d) sperma- 
theca strongly sclerotized and e) metathoracic wings: apical field with a vertical secon- 
dary sclerification near the radial cell. The genus Cyphopisthes can be divided into two 
groups, whose definition and status are being dealt with in a separate paper, Ebbritto- 
niella shares four further synapomorphies with the group formed by C. acromialis 


272 ALBERTO BALLERIO 


Fic. 5 — a: E. gestroi (Cameron Highlands), aedeagus with accessory sclerite (AS) (scale bar: 1 
mm); b: E. gestroi (Cameron Highlands), parameres (lateral view) (scale bar: 0,1 mm); c: 
E. gestroi (Cameron Highlands) genital segment (scale bar: 1 mm); d: E. gestroi (Cameron 
Highlands), bursa copulatrix with sclerite (S) and spermatophore inside (scale bar: 1 mm). 


REVISION OF THE GENUS EBBRITTONIELLA 273 


h 


Fic. 6 — a (Lata Jarom), b (Lata Jarom), c (Lata Jarom), d (Sabah: Mendolong): E. ignita, 
spermatheca; e (Borneo: north of Kalabakan), f (Maxwell Hill), g (Cameron Highlands), h 
(Cameron Highlands): E. gestroi, spermatheca (scale bar: 0,1 mm). 


274 ALBERTO BALLERIO 


(Pascoe, 1860) and few other species, i.e. f) labium with a very deep and wide regular- 
U-shaped excavation in the middle, g) spermatheca deeply U-shaped, h) bursa 
copulatrix with a characteristically shaped sclerite, and i) presence of an accessory 
sclerite among parameres. Characters b), c) and h) are autoapomorphic. 

The similarities with Ceratocanthus White, 1842 stressed by earlier authors are 
likely due to convergent evolution and, although at the present stage of knowledge 
remote relationships cannot be excluded, there are several characters that place 
Ebbrittoniella and Cyphopisthes quite far from Ceratocanthus. The latter differs at least 
in the following characters: a) labrum neither abruptly truncate nor divided by any 
carina, b) elytra without a false epipleuron, c) metathoracic wings: MP,,, — RP loop 
absent, d) metathoracic wings: apical field without vertical secondary sclerification, e) 
meso- and meta- tarsi capable of being folded along the longitudinal axis of the inner 
face of the tibia, f) sexual dimorphism not involving the shape of apical teeth of 
protibiae and g) spermatheca weakly sclerotized. 


ACKNOWLEDGEMENTS 


I wish to thank the following people for permission to study material, loan of 
specimens, valuable information and advice: S. Adebratt (Frinnaryd, Sweden), D. 
Ahrens (SMTD), S. Beévai (Ceské Budéjovice, Czech Republic), S. Bily (NMPC), Y. 
Cambefort (MNHN), P. Cechovsky (Brno, Czech Republic), G. Cuccodoro (MHNG), I. 
Jenis (Naklö, Czech Republic), M.D. Kerley (BMNH), J. Krikken (RMNH), I. Löbl 
(MHNG), O. Martin (ZMUC), G. McGavin (OXUM), O. Merkl (HNHM), O. 
Montreuil (MNHN), R. Paulian (Bordeaux), R. Poggi (MCSN), G. A. Samuelson 
(BPBM), C.H. Scholtz (University of Pretoria), J. Scheuern (Westum, Germany), H. 
Schönmann (NHMW), R. Schuh (Katzelsdorf, Austria), C. Zorn (Dresden). R. Furlan 
(Centro C.U.G.A.S., Universita di Padova) took SEM micrograph and J. Kobylak 
(Praha) executed the habitus drawings. 


REFERENCES 


Arrow, G. J. 1912. Coleopterorum Catalogus pars 43 - Scarabaeidae: Pachypodinae, Pleoco- 
minae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae, 
Dynamopinae, Acanthocerinae, Troginae. W. Junk, Berlin, 66 pp. 

BALLERIO, A. 1999. Revision of the genus Pterorthochaetes, first contribution (Coleoptera: 
Scarabaeoidea: Ceratocanthidae). Folia Heyrovskyana 7(5): 221-228. 

BALLERIO, A. 2000. A new genus and species of Ceratocanthidae from Tanzania (Coleoptera: 
Scarabaeoidea). African Zoology 35(1): 131-137. 

BROWNE, D. J. & SCHOLTZ, C. H. 1995. Phylogeny of the families of Scarabaeoidea (Coleoptera) 
based on characters of the hindwing articulation, hindwing base and wing venation. 
Systematic Entomology 20: 145-173. 

BROWNE, D. J. & SCHOLTZ, C. H. 1996. The morphology of the hind wing articulation and wing 
base of the Scarabaeoidea (Coleoptera) with some phylogenetic implications. Bonner 
zoologische Monographien 40: 1-200. 

BROWNE, J. & SCHOLTZ, C. H. 1998. Evolution of the scarab hindwing articulation and wing base: 
a contribution toward the phylogeny of the Scarabaeidae (Scarabaeoidea: Coleoptera). 
Systematic Entomology 23: 307-326. 


REVISION OF THE GENUS EBBRITTONIELLA 275 


D’HoTMan, D. & SCHOLTZ, C. H. 1990. Phylogenetic significance of the structure of the external 
male genitalia in the Scarabaeoidea (Coleoptera). Entomology Memoir Department of 
Agricultural Development Repubublic of South Africa 77,51 pp. 

GESTRO, R. 1899. Sopra alcune forme di Acanthocerini. Annali del Museo civico di Storia natu- 
rale di Genova XXXIX: 450-498. 

HOWDEN, H. F. & GILL, B. D. 1995. Trachycrusus, a new genus of Ceratocanthinae (Coleoptera 
Scarabaeidae) with two new species from Peru. The Canadian Entomologist 127: 
587-593. 

KUKALOVA-PECK, J. & LAWRENCE, J. F. 1993. Evolution of the hind wing in Coleoptera. The 
Canadian Entomologist 125: 181-258. 

LANSBERGE, J. W. VAN 1887. Trogides nouveaux. Notes from the Leyden Museum 1X:199-211. 

MARTINEZ, A. 1962. Un nuevo género de Acanthocerinae (Col. Scarabaeidae). Physis 23 n. 64: 
61-64. 

NEL, A. & DE VILLIERS, W. M. 1988. Mouthpart structure in Adult Scarab Beetles (Coleoptera 
Scarabaeoidea). Entomologia Generalis 13 (1/2):95-144. 

NEL, A. & SCHOLTZ, C. H. 1990. Comparative morphology of the mouthparts of adult Scara- 
baeoidea (Coleoptera). Entomology Memoir Department of Agricultural Development 
Repubublic of South Africa 80, 84 pp. 

PAULIAN, R. 1942. Coléoptéres Acanthocérides nouveaux ou peu connus. Revue francaise 
d’Entomologie IX: 70-75. 

PAULIAN, R. 1978. Revision des Ceratocanthidae [Col. Scarabaeoidea] II - Les espèces orientales 
et australiennes. Annales de la Societe entomologique de France (N.S.) 14 (3): 479-514. 

PREUDHOMME DE BORRE, A. 1886. Catalogue des Trogides décrits jusqu’à ce jour, précédé d’un 
synopsis de leur genres et d’une esquisse de leur distribution géographique. Annales de la 
Société entomologique de Belgique 30: 54-82. 

WESTWOOD, J. O. 1883. Two new species of the coleopterous genus Acanthocerus. Notes from 
the Leyden Museum 5: 1-2. 


iP 


î sd | ity I 
tari 


REVUE SUISSE DE ZOOLOGIE 107 (2): 277-281; juin 2000 


A new desert psocid from Namibia (Insecta: Psocoptera: Trogiidae) 


Charles LIENHARD 
Museum d’histoire naturelle, c. p. 6434, CH-1211 Geneve 6, Switzerland. 
E-mail: charles.lienhard @ mhn.ville-ge.ch 


A new desert psocid from Namibia (Insecta: Psocoptera: Trogiidae). - 
A single female of a new species representing a new genus of the family 
Trogiidae (Psocoptera: Trogiomorpha: Atropetae) is described and illus- 
trated: Spinatropos philippi gen. n., sp. n. The specimen has been collected 
in an unusual biotope for Psocoptera, at Diaz Point (Liideritz), on a rocky 
desert headland with hardly any vegetation. 


Key-words: Psocoptera - Trogiidae - new genus - new species - desert 
fauna - Namibia. 


INTRODUCTION 


Almost nothing is known on Psocoptera of Namibia. The only psocid recorded 
from this country (from Windhoek) is the cosmopolitan and usually domestic species 
Liposcelis bostrychophila Badonnel, 1931 (cf. Checklist of Southern African Insects on 
Internet: http://www.ru.ac.za/departments/zooento/Martin/Insects.html). 

The present material has been collected by N. P. & M. J. Ashmole under stones 
on a rocky desert headland with hardly any vegetation near Diaz Point (Liideritz), a 
very unusual biotope for Psocoptera. These insects generally live on vegetation, mostly 
trees or shrubs, or in leaf-litter (cf. Lienhard, 1998a). An important factor which enables 
a psocid to live in the above mentioned biotope could be the proximity to the sea, which 
guarantees a constantly high relative humidity of the air. The occurrence of a psocid of 
the genus Liposcelis Motschulsky, 1852 on apparently “sterile” littoral limestone rocks 
in the Mediterranean (Cyprus) has been reported by Lienhard (1998b). It would be 
worthwhile for ecologists to pay more attention to psocids accidentally captured in such 
unusual situations. 

Besides these ecological aspects the present material is also interesting from a 
taxonomic and perhaps ethological point of view. It represents a very characteristic new 
species of the family Trogiidae which cannot be placed in one of the hitherto known 
genera of this family. Its most striking diagnostic character, the presence of two short 
stout externoapical spurs on each of the somewhat elongated third valvulae (fig. 1) may 
prove to be connected with special ovipositional behaviour. As known up to now, all 
Psocoptera lay their eggs on the surface of a substrate (cf. Lienhard, 1998a). The new 
species, however, seems to be equipped to slightly dig in its eggs into a sandy soil. 


Manuscript accepted 18.01.2000 


278 CHARLES LIENHARD 


According to N. P. Ashmole (in litt.) there were small deposits of sand or fine rock 
fragments in the crevices of its habitat at Diaz Point. Similar digging spines on 
ovipositor are known in other insects, e.g. Diptera, where they are present in sand- 
dwelling Therevidae (cf. Irwin & Lyneborg, 1981: p. 514, fig. 22) or in the essentially 
dune-inhabiting species Helina protuberans Zett. (Muscidae), which lays its eggs into 
sandy soils (cf. Hennig, 1964: p. 209, Textfig. 46). 


Spinatropos gen. n. 


Diagnosis. Apterous, mesothorax in dorsal view about of same width as 
prothorax, metathorax distinctly wider. Body densely pubescent, excepting the almost 
bald postclypeus, which bears only a few peripheral hairs. Postclypeus of normal shape, 
not exceedingly bulging. Compound eyes well developed, hemispherical, with some 
small hairs in dorsal half, ocelli absent. Maxillary palpus (fig. 4) with fourth (terminal) 
segment (P4) of cylindrical shape, some stout setae present in apical half of P2, P3 and 
P4, no forked sensillum on P4. Laciniae symmetrically developed, lacinial tip 
tridentate, teeth diverging and well developed (fig. 3). Hind tibia with two apical spurs 
and two spurs on inner face in apical half. Pretarsal claw lacking preapical tooth, 
pulvillus long and slender, its tip slightly enlarged (fig. 6). 8th sternite of the female 
without sclerified knob which could be used in sound production (cf. Lienhard, 1998a: 
fig. 26m, Trogium pulsatorium). Third valvula (fig. 1) somewhat elongated, as typical 
of the family-group Atropetae, with spiny pilosity, two short stout externoapical spurs 
and one slender normal seta near tip. Second valvula (fig. 1) relatively well developed, 
reaching almost to the middle of the third valvula. Spermapore simple (fig. 5). 
Spermatheca with two oval parietal glands of about equal dimensions, situated close to 
each other in the proximal zone of the vesicle (fig. 7). Glands with numerous pores, no 
papillae (fig. 8). Spermatophore with very long tubular part (fig. 7). Male unknown. 

Type species. S. philippi sp. n. 

Etymology. Spina (lat.) = spine, an allusion to the particular pilosity of the third 
valvula. Atropos = one of the three Fates, the antique goddesses of destiny; Atropos 
Leach, 1815 is a synonym of Trogium Illiger, 1798 and gave rise to the family-group 
name Atropetae. In analogy to Atropos the new name is of feminine gender. 

Discussion. See discussion of the type species. 


Spinatropos philippi sp. n. Figs 1-8 


MATERIAL 

Holotype 9. Namibia: Diaz Point (Liideritz), rocky desert biotope with hardly 
any vegetation, relatively close to the sea, by visual seaching under stones, 30.X1.1994, 
leg. N. P. & M. J. Ashmole (sample nr 3090) (Muséum d'histoire naturelle de Genève). 


ETYMOLOGY 

The species is dedicated to N. Philip Ashmole, one of its collectors, in 
acknowledgement of his tireless efforts in collecting psocids in biotopes where they 
“should not occur”. 


A NEW DESERT PSOCID FROM NAMIBIA 279 


DESCRIPTION (2) 

Coloration. Body, legs and antenna whitish to light brown. Compound eye black 
(after 4 years in alcohol). Apical half of flagellar segments darker brown than basal half 
in basal part of the antenna, flagellar segments in its apical part entirely brown. 

Morphology. See generic diagnosis, with the following complements. Vertical 
and frontal sutures distinct. Both antennae damaged, one with 22, the other with 11 
segments. Marginal sensilla of labrum (fig. 2) typical of the family Trogiidae (cf. 
Badonnel, 1977). The complete aptery of the specimen is real and not due to broken 
winglets. This is supported by the absence of insertion points of winglets and by the 
even pilosity of thoracic tergites (in the frequently encountered cases of micropterous or 
brachypterous Trogiidae where winglets are broken, insertion points are visible on 
mesothoracic tergite and the lateral parts of the metathoracic tergite are bald, as it has 
been figured for the genus Cerobasis Kolbe, 1882 by Lienhard, 1998a: fig. 24c). 
Mesothorax (in dorsal view) shorter than half width of vertex. No coxal organ differen- 
tiated. Femora somewhat thickened, hind femur about three times as wide as hind tibia. 
Hind tibia with some long hairs on outer face. First segment of hind tarsus with 
numerous stout spur-like setae on ventral face. Epiproct bald in the middle, with some 
lateral hairs. Paraprocts with a distinct anal spine on hind margin. Subgenital plate 
absent, ovipositor valvulae basally covered by some membraneous folds. The wall of 
the spermathecal vesicle bears numerous fine pores, especially in the more distal half 
(fig. 7). The length of the oval spermathecal glands is about 125 um, each gland bears 
much more than 100 pores (fig. 8). 

Measurements. Body length = 2,0 mm. Vertex (width of head capsule) = 650 
um. Length of hind femur = 520 um. Length of hind tibia = 750 um. Length of hind 
tarsomeres (measured from condyle to condyle): tl = 320; t2 = 72; t3 = 90. 

Remark. The male is not yet known, but the presence of a spermatophore in the 
spermatheca of the female gives evidence of the bisexuality of the species. 


DISCUSSION 


Within the family Trogiidae (sensu Smithers, 1990) the new genus is charac- 
terized by the very particular spiny pilosity of the third valvula, especially by the 
presence of two stout externoapical spurs, but also by the cylindrical shape of the 
terminal segment (P4) of the maxillary palpus. In all other non-fossil genera, P4 is at 
least slightly enlarged apically, often much enlarged (hatchet-shaped). In the genera 
where females are known (not known in Anomocopeus Badonnel, 1967) the pilosity of 
the third valvula consists of normally shaped slender setae, as usual in the family-group 
Atropetae. The genus can also be distinguished from most of the other genera (amber 
fossils included) by its complete aptery. The only other apterous genera are Anomo- 
copeus and Mymicodipnella Enderlein, 1909; some apterous species are also known in 
the large genus Cerobasis. 

Nothing exact is known about the biology of the new species, for some general 
remarks, see the introduction. 


280 CHARLES LIENHARD 


Fics 1-8 


Spinatropos philippi gen. n., sp. n., female: 1, ovipositor valvulae; 2, marginal sensilla of labrum; 
3, lacinial tip: 4, maxillary palpus: 5, spermapore; 6, pretarsal claw: 7, spermatheca, containing 
one spermatophore: 8, parietal gland of spermatheca. 


A NEW DESERT PSOCID FROM NAMIBIA 281 


ACKNOWLEDGEMENTS 


I am very grateful to N. Philip and Myrtle J. Ashmole (Edinburgh) for having 
made their Namibian collections of psocids available to me. I also thank N. P. Ashmole 
for reading the manuscript and making some valuable suggestions. I am indebted to my 
colleague Bernhard Merz (Geneva) for information about the presence of digging 
spines on the ovipositor in Diptera. 


REFERENCES 


BADONNEL, A. 1977. Sur les sensilles du bord distal du labre des Psocoptères et leur intérêt 
taxonomique. Bulletin de la Société entomologique de France 82(5-6): 105-113. 

HENNIG, W. 1964. Muscidae (pp. 1-624). In: LINDNER, E. (ed.). Die Fliegen der palaearktischen 
Region. Vol. 7(2), part 63b(1). 

IRWIN, M. E. & LYNEBORG, L. 1981. Therevidae (pp. 513-523). In: Manual of Nearctic Diptera. 
Vol. 1. Research Branch Agriculture Canada, Monograph 27: VI + 674 pp. 

LIENHARD, C. 1998a. Psocopteres euro-méditerranéens. Faune de France 83: XX + 517 pp. 

LIENHARD, C. 1998b. Interessantes aus der Welt der Psocopteren. Mitteilungen der Schwei- 
zerischen Entomologischen Gesellschaft 71(1-2): 252-253. 

SMITHERS, C. N. 1990. Keys to the families and genera of Psocoptera (Arthropoda: Insecta). 
Technical Reports of the Australian Museum 2: 1-82. 


Eu 


L 


REVUE SUISSE DE ZOOLOGIE 107 (2): 283-323; juin 2000 


Terrestrial Isopoda from Guatemala and Mexico 
(Crustacea: Oniscidea: Crinocheta) 


Andreas LEISTIKOW 

Universität Bielefeld 

Abteilung für Zoomorphologie und Systematik 
Morgenbreede 45 

D-33615 Bielefeld. 

E-mail: Leiste@Biologie.Uni-Bielefeld.de 


Ruhr-Universität Bochum 
Lehrstuhl für spezielle Zoologie 
Universitätsstraße 150 
D-44780 Bochum. 


Terrestrial Isopoda from Guatemala and Mexico (Crustacea: Onisci- 
dea: Crinocheta). - Some terrestrial Isopoda from Mexico, described in 
the genus Philoscia Latreille, 1804, are redescribed and transferred to other 
genera. Two new genera, Quintanoscia gen. n. and Oxalaniscus gen. n. are 
described, they belong to the most primitive representatives of the Onis- 
coidea within the Crinocheta. One of the species, Androdeloscia formosa 
(Mulaik, 1960) is recorded from Guatemala for the first time. A closely 
related species, A. valdezi sp. n. is described as new to science. New 
records for Littorophiloscia denticulata (Ferrara & Taiti, 1981) and Burmo- 
niscus kohleri (Schmalfuss & Ferrara, 1978) are presented; they were 
recorded in the New World for the first time. Their distributional patterns 
and phylogenetic relationships are discussed. 


Key-words: Crustacea - Oniscidea - Philoscia - Androdeloscia - Littoro- 
philoscia - Burmoniscus - new genera - new species - Central America - 
taxonomy. 


INTRODUCTION 


Central America is one of the regions of the world, which is almost unexplored 


with respect to the terrestrial isopod fauna. Only few contributions deal with taxa of 
Oniscoidea collected in this biogeographically interesting area, as e.g., Miers (1877) 
reporting on some Oniscidea from Central America. Particularly isopods of the 
northern countries, Mexico and Guatemala, are little known. Richardson (1907) 
described a new scleropactid species Spherarmadillo schwarzi Richardson, 1907, 
from Guatemala and later a species of Armadillidae, Globarmadillo armatus Richard- 


a Manuscript accepted 05.01.2000 


284 ANDREAS LEISTIKOW 


son, 1910 was described by Richardson (1910). This work remains the only contri- 
bution to the Guatemalan oniscidean fauna. 

The exhaustive contribution of Mulaik (1960) on the terrestrial isopod fauna of 
Mexico is an important work, although the taxonomy used was no more up to date in 
1960. Many species placed in the genus Philoscia Latreille, 1960 do not show any 
apomorphic character in common with, e.g., the European Philoscia muscorum 
(Scopoli, 1793), which is a typical member of this genus. Some of the species 
included in Philoscia are among the most primitive representatives of Oniscoidea as 
defined by Schmalfuss (1989): they, for instance, lack the noduli laterales. Some 
species of Philoscia were revised in this study and transferred to other genera. 

More recently, Schultz (1977) contributed to our knowledge of the troglobitic 
species of Oniscidea from Central America, with Troglophiloscia laevis Schultz, 1977 
being added to the Mexican fauna. The species is related to the Cuban 7. silvestrii 
Brian, 1929, the type of the genus (Brian, 1929). The species might also occur in 
northern Guatemala, as there are many caves in the northern province Petén, which 
might be suitable habitats for this genus. However, it was not found in the Cueva 
Actun Can near Santa Elena, Petén in a recently started survey. 

This study aims to increase our knowlegde on the crinochete Oniscidea , which 
are of philosciid appearance. Several species are described in detail to get access to a 
complete data set for a phylogenetic analysis. New data are given for some species 
probably introduced to Guatemala. Both Littorophiloscia denticulata (Ferrara & Taiti, 
1981) and Burmoniscus kohleri (Schmalfuss & Ferrara, 1978) are recorded for the 
first time from the New World. Ther distribtional patterns and phylogenetic relation- 
ships are discussed. 

The material examined is deposited in: Instituto Politécnico Nacional de 
México (IPNM), Staatliches Museum für naturkunde (SMNS), Muséum d'histoire 
naturelle de Geneve (MNHG), Universidad del Valle de Guatemala (UVG) and in the 
author’s collection. 


SPECIES ACCOUNT 
Quintanoscia gen. n. 


DIAGNOSIS: Cephalothorax without linea frontalis and linea supra-antennalis; 
lamina frontalis present; compound eyes composed of 12 ommatidia; antennula 
composed of three cylindrical articles; antennal flagellum three-articulate with apical 
organ bearing two short free sensilla. 

Mandibles with molar penicil consisting of three branches; medial endite of 
maxillula with two penicils and apical tip: lateral endite with 4+6 teeth and slender 
stalk; lobes of maxilla subrectangular: densely covered with trichiae; maxilliped 
basipodite without sulcus lateralis; palp three-articulate; proximal article with two 
long setae; medial and distal article with prominent setal tufts; endite elongate; setose; 
with long penicil rostrally. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 285 


Pereopods stout; antenna-grooming brush of carpus | arranged longitudinally; 
dactylus with long inner claw; dactylar seta ending in a knob; coxal plates without 
sulcus marginalis and gland pores; noduli laterales present; quite similar to tricorn- 
like setae; insertion on all coxal plates at same distance from lateral margin. 

Pleopods with rhomboidal exopodites bearing sensory spines laterally: 
endopodites subquadrangular; no respiratory areas discernible at 400x magnification: 
male genital papilla with ventral shield. 

Uropod with protopodite laterally grooved; exopodite twice as long as the 
more proximally inserted endopodite. 


TYPE SPECIES: Philoscia contoyensis Mulaik, 1960 (by monotypy). 

NUMBER OF NOMINAL SPECIES: only type species. 

ETYMoLoGY: The genus is named after the Mexican province Quintana Roo, where 
the type species was collected. 

DISTRIBUTION: Only known from southeastern Mexico on the Yucatan Peninsula. 
REMARKS: The genus Quintanoscia gen. n. is close to the groundpattern of the 
Oniscoidea, the shape of the maxilliped is similar to Dero Guérin, 1836 and Allo- 
niscus Dana, 1852. The pereopod 1 is equipped with a longitudinal carpal brush and 
the medial margin is bearing several bifid sensory spines. The autapomorphies of the 
genus are: 


BM Reduction of the lateral lobes [lateral lobes present] 


m Cephalothorax broadened [vertex not broader than height of cephalothorax] 

The coxal plates bear several tricorn-like setae. One of those setae bears a 
distinctly longer sensillum, which is twice as long as the basal cuticular plaque. This 
structure can be interpreted as a nodulus lateralis. 


Quintanoscia contoyensis (Mulaik, 1960) Figs 1-6 


Philoscia contoyensis Mulaik, 1960 

Material: Paratypes, 5 d (max. body length 3.5 mm): Mexico, Quintana Roo, Isla 
Contoy, leg. 20.X1.1947, B.F. Osorio Tafall, IPNM 1631-A. 

Colour: Mulaik (1960) wrote in the original description: “La coloracion de los 
ejemplares conservados en alcohol; es un moteado de café; rojizo y armadillo. El 
color mas obscuro esta confiando a una franja dorsal; media; delgada e irregular; a 
una area angosta a lo largo de los margenes de os epimeros y entre estas; a otra banda 
mas angosta fragmentada en manchas. La region ventral es casi blanca. Las formes 
jovenes muestran menos pigmento que los adultos.” 

Cephalothorax: Rather large with regard to body length; compound eyes 
bearing 12 ommatidia; vertex strongly arched; no linea frontalis and linea supra- 
antennalis visible; lamina frontalis inconspicuous (fig. 1; Ctf). 

Pereon: Tegument smooth and shiny; bearing evenly spread tricorn-like setae; 
noduli laterales similar to tricorn-like setae; sensillum comparatively longer; on all 
coxal plates insertion at same distance to lateral margin; more distally located from 
plate I to VII; no sulcus marginalis and gland pores. 


286 ANDREAS LEISTIKOW 


Pleon: Rather short; narrower than pereon; prominent neopleurae on pleonites 
III to IV; pleotelson half as long than pleon; triangular; bordered by tricorn-like setae. 

Appendages: 

Antennula: Three-articulate with cylindrical articles; distal article bearing 
longitudinally arranged aesthetascs (fig. 1; Anl). 

Antenna: Peduncle covered with tricorn-like setae; flagellum three-articulate; 
medial and distal article with pair of aesthetascs; apical organ as long as medial 
article; with short free sensilla (fig. 5; An2). 

Mouth parts as described in generic diagnosis (fig. 2). 

Pereopods: Rather stout; carpus | with longitudinal antenna-grooming brush; 
medial border of carpus and merus 1 to 4 bearing many bifid sensory spines (fig. 3-5; 
PEI-7); dactylus with medium-sized inner claw (fig. 3; Dac); stout interungual seta; 
tricorn-like seta laterally; dactylar seta with knob-like tip (fig. 4; Sd5). Sexual 
dimorphism: Due to the lack of females not observed. 

Pleopods: Exopodites rhomboidal with lateral margin bearing four to ten 
sensory spines; on pleopod 4 and 5 one subapically on medial border; pleopod 5 with 
small pectinate scales caudally; not arranged in rows; endopodites rounded triangular 
to quadrangular; respiratory areas not discernible at 400x magnification (fig. 6; PLI- 
5). Sexual dimorphism: Pleopod | endopodite long; rather prominent with ill-defined 
basal area containing the intrinsic endopodite | levering muscle M49 (Erhard, 1997); 
no row of small spines medio-caudally (fig. 6; PL1); pleopod 2 exopodite with lateral 
margin slightly more sinuous than on exopodite 3; endopodite twice as long as 
exopodite; rather stout (fig. 6; PL2). 

Uropod: Protopodite triangular with lateral groove; endopodite inserting more 
proximally than two times longer exopodite (fig. 5; UR). 

Genital papilla: Ventral shield with parallel margins in basal half; slightly 
surpassed by orifices (fig. 6; Gen). 


Oxalaniscus gen. n. 


DIAGNOSIS: Cephalothorax with lamina frontalis; linea supra-antennalis and 
lateral lobes; linea frontalis reduced; compound eyes composed of about nine omma- 
tidia. Antennula three-articulate; slender; antenna with three-articulate flagellum 
(Mulaik, 1960). 

Mandible with molar penicil composed of three branches; lateral endite of 
maxillula with 4+5 simple teeth; slender stalk present; maxilla subrectangular; 
maxilliped with long penicil on endite and prominent setal tufts on palp. 

Pereopods with coxal plates lacking distinct nodulus lateralis; carpus 1 with 
longitudinal carpal brush; setal brushes composed of sensory spines; not very dense; 
present on carpus and merus | to 5; dactylar seta apically spatuliform. 

Pleopods with prominent exopodites bearing few sensory spines laterally; no 
respiratory structures discernible. 

Uropod with protopodite subtriangular carrying lateral groove; endopodite 
inserting slightly proximally of exopodite. Genital papilla with ventral shield sur- 
passed by terminal spatula. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 287 


/ 
EE —— 4 
100um / li 


Fic. 1: Quintanoscia contoyensis (Mulaik, 1960), 4 3.5mm body length. Anl antennula; Ctf 
cephalothorax in frontal view; Cx4 coxal plate IV; Had habitus in dorsal view; Hal habitus in 
lateral view; N14/7 setae of nodulus lateralis shape from coxal Figures IV and VII; Sx4 tricorn- 
like seta of coxal plate IV; Tel pleotelson. 


288 ANDREAS LEISTIKOW 


SOum 


Fic. 2: Quintanoscia contoyensis (Mulaik, 1960), 4 3.5mm body length. Mdl/r left and right 
mandible; Mxp maxilliped, with detail of endite in rostral view; Mx1 maxillula, with details of 
apical lateral endite in caudal and rostral view; Mx2 maxilla. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 289 


100um 


FIG. 3: Quintanoscia contoyensis (Mulaik, 1960), 3 3.5mm body length. Dac dactylus of pere- 
opod I in rostral view; PE1-4 pereopods | to 4, in rostral (PE1) or caudal (PE2-4) view; Sbl 
tricorn-like seta of basis 1; Scl sensory spine of carpal brush; Sc2 sensory spine of carpus 2. 


290 ANDREAS LEISTIKOW 


FIG. 4: Quintanoscia contoyensis (Mulaik, 1960), d 3.5mm body length. PES-6 pereopods 5 
and 6 in caudal view; Sb6 tricorn-like seta of basis 6; Sc5/7 sensory spines of carpus 5 and 7; 
Sd5 dactylar seta of dactylus 5. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 29] 


FIG. 5: Quintanoscia contoyensis (Mulaik, 1960), d 3.5mm body length. An2 antenna, with 
detail of apical organ; PE7 pereopod 7 in caudal view; UR uropod in rostral view. 


ANDREAS LEISTIKOW 


i) 
Ne} 
tO 


FIG. 6: Quintanoscia contoyensis (Mulaik, 1960), 4 3.5mm body length. Gen genital papilla: 
PL1-5 pleopods 1 to 5 in rostral view, with detail of endopodite 1 in caudal view. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 293 


TYPE SPECIES: Philoscia ctenoscoides Mulaik, 1960 (by monotypy). 
NUMBER OF NOMINAL SPECIES: Only type species included. 
ETYMOLOGY: The genus name is derived from oxalä, the god of creation of the 
indigenas from Brazil. 
REMARKS: The genus Oxalaniscus gen. n. belongs to the most basal species of the 
taxon Oniscoidea. It seems closely related to Quintanoscia gen. n. The putative 
synapomorphies are given above. The following characters are the autapomorphies of 
this genus: 

BI Slender stalk of maxillula in a more lateral position [slender stalk medially, 

surrounded by group of lateral teeth] 


In all other taxa with a slender stalk on the maxillula, it is in a medial position 
on the apical region, surrounded by the medial teeth of the outer group. 


Oxalaniscus ctenoscoides (Mulaik, 1960) Figs 7-10 


Philoscia ctenoscoides Mulaik, 1960 

Material: several specimens: Mexico, Chiapas, Ruinas de Palenque, leg. 13.VII.1949, 
C. and M. Goodnight, Finca Guautimoc, leg. 02.VII.1950, C. and M. Goodnight; Tabasco, 
Emiliano Zapata, leg. 15.VIII.1945; Quintana Roo, Puerto Morelos, leg. XI.1947, B.F. Osorio 
Tafall, Isla Cozumel, leg. 24.X1.1947, B.F. Osorio Tafall, IPNM 1628. 

Colour: “Dorso brillante e intensamente moteado con café” (Mulaik, 1960). 

Cephalothorax: Wider than high with linea supra-antennalis and lamina fron- 
talis; small lateral lobes present; compound eyes composed of about nine ommatidia 
(1 75 Cth): 

Pereon: Tegument smooth with few tricorn-like setae; coxal plates without 
gland pores; sulcus marginalis present; no noduli laterales visible (fig. 7, Cx4). 

Pleon: Only slightly narrower than pereon; with prominent neopleurae on 
pleonite 3 to 5; pleotelson with rounded apex; lateral margins straight. 

Appendages: 

Antennula: Slender; composed of three articles; distal one cylindrical with 
apical pair of aesthetascs and medial set of about four aesthetascs (fig. 7, Anl). 

Antenna: Peduncle with scattered tricorn-like setae; flagellum broken in all the 
material examined; thus nothing can be said about the shape and the apical organ. 
According to Mulaik (1960) it is composed of three articles. 

Mouth parts as decribed in generic diagnosis (fig. 8). 

Pereopods: Rather stout (fig. 9; PEI-7); carpus without latero-distal setal tuft; 
no ornamental sensory spine on carpus |; antenna-grooming brush longitudinal; 
carpus and merus | to 5 with loose brush of sensory spines; dactylus with short inner 
claw (fig. 9, Dac); all interungual setae broken in material examined; dactylar seta 
with flattened spatuliform apex (fig. 9, Sd1). 

Pleopods: Exopodites of pleopod 3 to 5 large; bearing one to two sensory 
spines laterally; no respiratory areas discernible in light microscope; endopodites 
subrectangular (fig. 10, PLI-5). Sexual dimorphism: Male pleopod 1 exopodite 
obtusely triangular; endopodite more than two times longer than exopodite; slightly 


294 ANDREAS LEISTIKOW 


bent sidewards; apex strongly bent sidewards with subterminal protrusions; about five 
to ten small spines at proximal end of spermatic channel (fig. 10, PL1). Pleopod 2 
exopodite triangular with single latero-distal sensory spine; distal third of endopodite 
lace-shaped; slightly bent (fig. 10, PL2). 

Uropod: As described in generic diagnosis. 

Genital papilla: Ventral shield surpassed by terminally ending orifices; apex 
indistinctly truncate (fig. 10, Gen). 
REMARKS: The two new genera Quintanoscia and Oxalanoiscus from Mexico are 
quite primitive with respect to several characters: They bear a subrectangular maxilla, 
the maxillipedal palp is equipped with very prominent setal tufts and the penicil of the 
endite is still prominent. Nonetheless, they are united in an sistergroup relationship 
due to the following synapomorphies: 


@ Cephalothorax with linea frontalis reduced [linea frontalis present] 

@ Molar penicil composed of three branches [molar penixcil composed of 
about ten branches] 

@ Lateral endite of maxillula with 6+4 teeth, one of the inner set absent or at 
least vestigial [no tooth reduced in size] 

@ Maxillular teeth simple [teeth of inner set cleft] 


In the outgroup, represented by Ligia baudiniana Milne Edwards, 1840, Deto 
echinata Guérin, 1836 several species of Scleropatidae and Alloniscus Dana, 1852, 
the characters differ considerably. The mandible is bearing a penicil of about ten 
unfused branches and the maxillular teeth of the inner set are cleft or ctenate. 
Therefore, the character states in both new genera are interpreted as being derived. 

In spite of their close relationship, each species is placed in a monotypic genus 
since the structure of the dorsal tricorn-like setae and the cephalothorax are striking. 
Probably Quintanoscia and Oxalaniscus are as distant to each other as they are to 
Alloniscus. In recent works on terrestrial isopods, the following characters are mostly 
used for generic separation: cephalothorax morphology, structure and position of 
dorsal receptors, presence of a sulcus marginalis, structure of the maxilliped, 
arrangement of the teth on the lateral endite of the maxillula, setal patterns of the 
pereopods, shape of pleon and pleotelson, shape of the male pleopod 5 and genital 
papilla (Vandel, 1973a, 1973b; Taiti & Ferrara, 1980). Therefore, a generic separation 
of the two taxa seems plausible. 


Genus Littorophiloscia Hatch, 1947 


DrAGNosis: Cephalothorax with linea supra-antennalis; slight lamina frontalis 
and lateral lobes; linea frontalis lacking; compound eyes composed of about 15 
ommatidia. Antennula three-articulate; slender; antenna long with three-articulate 
flagellum; apical organ longer than distal article. 

Mandible with molar penicil consisting of about ten free branches; maxillula 
with lateral endite bearing 4+6 teeth; inner set with five teeth cleft; slender stalk pre- 
sent; maxilla with both lobes subequal in width: maxilliped with endite setose; bea- 
ring prominent penicil rostrally; palp three-articulate; with three prominent setal tufts. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 295 


Cx4 


Fic. 7: Oxalaniscus ctenoscoides (Mulaik, 1960), d 3mm body length. Anl antennula; Ctf 
cephalothorax in frontal view; Cx4 coxal plate IV; Had habitus in dorsal view; Hal habitus in 
lateral view; Ste tricorn-like seta of pleotelson; Tel pleotelson. 


296 ANDREAS LEISTIKOW 


Fic. 8: Oxalaniscus ctenoscoides (Mulaik, 1960), d 3mm body length. Mdl/r left and right 
mandible; Mxp maxilliped, with detail of endite in rostral view; Mx1 maxillula, with detail of 
apical lateral endite in rostral view; Mx2 maxilla. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 297 


Na or LA 
HERE Cares (EA) 

GAL 7 \ 

Y & 

STA 
ge fy 
| > 4 
/ 


——— 
SOpim 


Fic. 9: Oxalaniscus ctenoscoides (Mulaik, 1960), 4 3mm body length. Dac dactylus of 
pereopod 1; PE1-7 pereopods | to 7 in rostral (PE1) or caudal view; Scl sensory spine of carpal 
brush; Sd1 dactylar seta of pereopod 1. 


298 ANDREAS LEISTIKOW 


Fic. 10: Oxalaniscus ctenoscoides (Mulaik, 1960), ¢ 3mm body length. Gen genital papilla; 
PL1-5 pleopods 1 to 5 in rostral view, with detail of endopodite 1 in caudal and rostral view. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 299 


Pereopods slender; ornamental sensory spine of carpus 1 with serrate double- 
fringe; dactylus with interungual seta long; more conspicuous than dactylar seta; inner 
claw short; coxal plates without gland pores; sulcus marginalis and nodulus lateralis 
present; all noduli inserted at same distance from lateral margin. Male pereopod 1 
propus and carpus inflated; bearing more or less prominent setal brushes. 

Pleopods without respiratory areas; exopodites with about five sensory spines 
laterally; endopodites bilobate. Uropod with protopodite laterally grooved; endopodite 
inserting more proximally than exopodite. 

Genital papilla with short ventral shield; orifices on small to conspicuously 
elongate protrusions of the terminal spatula. 


TYPE SPECIES: Philoscia richardsonae Holmes & Gay, 1904. 

NUMBER OF NOMINAL SPECIES: 23, four known from the Neotropics. 

REMARKS: Taiti & Ferrara (1986a) revised the genus and gave a diagnosis for it, they 
stressed the similarity of the male pereopods 1 and the mouth parts in Littorophiloscia 
and Halophiloscia Verhoeff, 1908. With respect to the mouth parts, the polatity of the 
characters was misinterpreted; they are of a plesiomorphic character state in both 
genera, similar to the above described genera Quintanoscia and Oxalaniscus, and to 
other Crinocheta with a position basal to the Oniscoidea sensu Schmalfuss (1989). 


Littorophiloscia denticulata (Ferrara & Taiti, 1981) Figs 11-14 


Bilawrencia denticulata Ferrara & Taiti, 1981 

Material examined: 3 4,9 2 (ovigerous), 3 juveniles: Guatemala, Izabal, Livinston, 
beach north of mouth of Rio Queguéche, between shells and fibres of coconuts, under Ipomoea 
sp., leg. 22.X.1998, A. Leistikow, MNHG and author's collection. 

Colour: Dorsally light violaceous brown with white paramedian patches; coxal 
plates with two to three white patches; cephalothorax densely spotted with white; 
pleon uniformly violaceous brown. 

Cephalothorax: Linea frontalis reduced; linea supra-antennalis; lamina fron- 
talis and small lateral lobes present; compound eyes comprising eight ommatidia (fig. 
BIAC): 

Pereon: Slightly arched; coxal plates with sulcus marginalis and nodulus 
lateralis; distance between noduli and lateral margin subequal on all coxal plates (fig. 
11, Cx4/Cxp); scattered tricorn-like setae present. 

Pleon: Narrower than pereon; small neopleurae on pleonites 3 to 5 present; 
pleotelson rounded with few tricorn-like setae along margin. 

Appendages: 

Antennula: Long and slender; three-articulate with pairs of aesthetascs in a 
rowon distal article (fig. 11, Anl). 

Antenna: Slender sparsely covered with of tricorn-like setae; flagellum three- 
articulate with long apical organ; bearing short free sensilla; being as long as distal 
article (fig. 11, An2). 

Mandible: Pars intermedia with two penicils on left and one on right side; 
bearing few setae; molar penicil composed of three branches; mandibular body 
slender (fig. 12, Mdl/r). 


300 ANDREAS LEISTIKOW 


Maxillula: Medial endite with two weak penicils and apical tip; lateral endite 
with 4+6 teeth; five of inner set cleft; slender stalk present (fig. 12, Mx1). 

Maxilla: Subrectangular with both lobes subequal in width; scattered 
trichiform setae present; more than ten cusps apically of medial lobe (fig. 12, Mx2). 

Maxilliped: Basipodite slender; with sulcus lateralis; palp three-articulate; setal 
tufts prominent; setae of proximal article subequal in length; endite slender; apically 
setose; prominent penicil on rostral surface present (fig. 12, Mxp). 

Pereopods: Slender; with scattered tricorn-like setae (fig. 13, PE1-7); carpus 
with prominent antenna-grooming brush; along medial margin covering almost one 
half of carpus length; ornamental sensory spine with hyaline apex; slightly striate (fig. 
13, Scl); dactylus with simple dactylar seta (fig. 13, Dac); inner claw short; inter- 
ungual seta with hyaline spatuliform apex. Sexual dimorphism: Male pereopods | to 2 
with setal brushes on carpus; propus | inflated; equipped with prominent setal brush. 

Pleopods: Pleopod exopodites rhomboid; with one to two sensory spines 
laterally; endopodites subtriangular; no respiratory organs discernible in light micro- 
scope (fig. 14, PLI-5). Sexual dimorphism: Male pleopod 1 endopodite rounded; 
endopodite slender; more than two times longer than exopodite; apex acute; slightly 
bent sidewards; distal third with caudomedial row of spines; laterally with three to 
five subapical teeth (fig. 14, PL1). Pleopod 2 exopodite pointed; laterally sinuous; 
endopodite slender; one third longer than exopodite (fig. 14, PL2). 

Uropod: As in other species of the genus. 

Genital papilla: Ventral shield with straight margins basally; terminal spatula 
with orifices surpassing ventral shield; slightly bilobate (fig. 14, Gen). 

DISTRIBUTION: Andaman Islands, Grub Island. 

REMARKS: Littorophiloscia denticulata was described from the Andaman Islands as a 
close relative of L. albicincta (Vandel, 1973) and L. occidentalis (Ferrara & Taiti, 
1983). The outstanding autapomorphy of this species is the denticulated male pleopod 
| endopodite; which is unique among its congeners. On the other hand, the species 
mentioned above have a pleopod | with a similar structure. It is not possible to 
evaluate the phylogenetic status of this character. Due to the relative simplicity of its 
structure in comparison to its congeners, like L. richardsonae (Holmes and Gay, 
1904) and L. riedlii (Strouhal, 1966), the latter formerly placed in Stenophiloscia 
(Strouhal, 1966), L. denticulata may be close to the basis of this genus. 

The occurence of this species in Guatemala most probably is due to human 
introduction. The locality is 20 km north of Puerto Barrios, the main sea harbour of 
Guatemala. L. denticulata may have been displaced with cargo from the South East 
Asia, where it is assumed to be autochthonous, because its closest relatives occur in 
the Indopacific region (Taiti & Ferrara, 1986a) and the radiation of this group might 
have taken place in this area. 


Burmoniscus Collinge, 1914 


A diagnosis of Burmoniscus was given in Taiti & Ferrara (1986b). 
TYPE SPECIES: Burmoniscus moulmeinus Collinge, 1914 (senior synonym of 
Philoscia coeca Budde-Lund, 1895). 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 301 


250um 


"n 
100um 


Fic. 11: Littorophiloscia denticulata (Ferrara & Taiti, 1981), 4 3.5mm body length. Anl 
antennula; An 2 antenna, with detail of apical organ; Ctf cephalothorax in frontal view; Cx4 
coxal plate IV, with detail of nodulus lateralis; Cxp coxal plates with position of noduli 


laterales: Had habitus in dorsal view; Hal habitus in lateral view; Ste tricorn-like seta of 
pleotelson; Tel pleotelson. 


302 ANDREAS LEISTIKOW 


| 
Fic. 12: Littorophiloscia denticulata (Ferrara & Taiti, 1981), ¢ 3.5mm body length. Mdl/r left ° 
and right mandible; Mxp maxilliped, with detail of endite in rostral view; Mx! maxillula, with 
details of apical lateral endite in rostral view; Mx2 maxilla. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 303 


Fic. 13: Littorophiloscia denticulata (Ferrara & Taiti, 1981), d 3.5mm body length. Dac 
dactylus of pereopod 1 in rostral view; PE1-7 pereopods 1 to 7 in caudal or rostral (PE1) view; 
Sb4 tricorn-like seta of basis 4; Scl ornamental sensory spine of carpus 1. 


304 ANDREAS LEISTIKOW 


100um 


RISI 


___ 


I OOum — PL4 


Fic. 14: Littorophiloscia denticulata (Ferrara & Taiti, 1981), 4 3.5mm body length /2 4.5mm 
body length. Gen genital papilla; PEf female pereopod | in rostral view; PL1-5 pleopods 1 to 5 
in rostral view, with detail of endopodite 1 in caudal view. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 305 


NUMBER OF NOMINAL SPECIES: 58, two are reported from the Neotropics, both 
of which most pobably introduced from West Africa and Southeast Asia. 

DISTRIBUTION: Southeast Asia, East Africa, West Africa (?); some species with 
circumtropical distribution due to dispersal by man. 

REMARKS: The genus Burmoniscus Collinge, 1914 has been revised and newly 
defined by Taiti & Ferrara (1986b). It is a heterogeneous, probably paraphyletic 
group, comprising blind and unpigmented species and species which are superficially 
similar to several members of /schioscia Verhoeff, 1928; e.g., Burmoniscus davisi 
Taiti & Manicastri, 1988. In fact, no autapomorphies of the genus are described until 
now. The characteristic position of the nodulus lateralis on coxal plate II 1s shared 
with, e.g., Anchiphiloscia Stebbing; 1908 (Ferrara & Taiti, 1986), from which it 
differs in the presence of gland pores and a sulcus marginalis, certainly plesio- 
morphies of Burmoniscus. 

The genus is distributed in the Oriental region, where it occurs with more than 
30 species. The first record from South America was B. meussii (Holthuis, 1949), 
reported from Brazil by Aratjo et al. (1996). It is found in suitable habitat in 
Venezuela as well (pers. obs.). Records of the little known species Burmoniscus 
kohleri (Schmalfuss & Ferrara, 1978) are presented from Guatemala for the first time. 


Burmoniscus kohleri (Schmalfuss & Ferrara, 1978) Figs 15-18 


Rennelloscia kohleri Schmalfuss & Ferrara, 1978 


Material examined: 2 d (max.body length 2.5 mm), several 9: Guatemala, Izabal, 
Aldea de Los Irrayoles, in scarse leaf litter in a plantation of Biepheridium? guatemalensis 
(Rubiaceae), leg. A. Leistikow; 17.X.1998, MNHG and author's collection; 2 4 (max. body 
length 2.5 mm); 5 9: Guatemala, Izabal, Aldea de Los Irrayoles; in a meadow with Cyperus 
sp.; leg. 17.X.1998, A. Leistikow, author's collection; 2 4; 3 2: Guatemala, Izabal, Livinston, 
finca in the western part of town on hill, under rotting stems of Cocos palms, leg. A. Leistikow; 
21.X.1998; author's collection 


Colour: Violaceous brown with light patches in medial line; paramedially and 
on coxal plates; vertex densely spotted with light patches; pleon uniformly violaceous 
brown. 

Cephalothorax: Vertex slightly arched; compound eyes composed of about five 
ommatidia; no linea frontalis and lamina frontalis; linea supra-antennalis present (fig. 
1155 (Caop 

Pereon: Slender; tegument smooth and shiny; scattered tricorn-like setae; coxal 
plates with sulcus marginalis and nodulus lateralis; noduli of coxal plate II and IV 
more remote from margin (fig. 15, Cx3/Cxp). 

Pleon: Narrower than pereon; no neopleurae visible in dorsal view; pleotelson 
with sinuous lateral margins; pointed; bearing few tricorn-like setae. 

Appendages: 

Antennula: Proximal article wide; medial and distal article forming a cone; 
aesthetascs at top of distal article (fig. 15, Anl). 

Antenna: Peduncle five-articulate; length ratio from proximal to distal joint 1: 
2: 2: 3: 4; flagellum three-articulate with pairs of aesthetascs on medial and distal 
article; apical organ as long as distal article; with short free sensilla (fig. 15, An2). 


306 ANDREAS LEISTIKOW 


SOum 


Fic. 15: Burmoniscus kohleri (Schmalfuss & Ferrara, 1978), ¢ 2.7mm body length. Anl 
antennula; An 2 antenna, with detail of apical organ; Ctf cephalothorax in frontal view; Cx3 
coxal plate III; Cxp coxal plates with position of noduli laterales; Had habitus in dorsal view; 
Hal habitus in lateral view; Tel pleotelson. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 307 


Mandible: Molar penicil simple; pars intermedia with sparse setation; two 
penicils on left; one on right side; intermedial penicil present (fig. 16, Mdl/r). 

Maxillula: Medial endite pointed; bearing two penicils apically; lateral endite 
with 4+5 teeth; three of inner set cleft; one double-cleft; one simple (fig. 16, Mx1). 

Maxilla: Lateral lobe more than two times broader than medial one; setose; 
laterally with transverse setal rows; medial lobe with few setae and five apical cusps 
(fig. 16, Mx2). 

Maxilliped: Basipodite with sulcus lateralis; palp with proximal article bearing 
two setae; inserted close together; distal articles fused; two setal tufts present; pro- 
ximal one with one prominent seta; endite with strong tooth caudally and knob-like 
penicil rostrally (fig. 16, Mxp). 

Pereopods: Slender with latero-distal setal tuft on carpus (fig. 17, PE1-7); 
prominent transverse setal brush on rostral surface of carpus 1; half as broad as 
medial margin; ornamental sensory spine double fringed (fig. 17, Sc1); dactylus with 
short inner claw (fig. 17, Dac); dactylar seta simple; rather short. No sexual dimor- 
phism discernible. 

Pleopods: Pleopod exopodites rhomboid; bearing one to two sensory spines 
laterally; single row of pectinate scales on caudal surface of exopodite 5; no 
respiratory areas discernible; endopodites bilobate (fig. 18, PL1-5). Sexual dimor- 
phism: Male pleopod 1 exopodite strongly concave on lateral margin; sinuosity 
forming almost right angle; endopodite bent laterally; tapering in distal third; medio- 
caudal row of short spines present (fig. 18, PL1). Pleopod 2 exopodite pointed; 
laterally sinuous; endopodite slender (fig. 18, PL2). 

Uropod: As in other species of the genus. 

Genital papilla: Ventral shield stout; slightly surpassed by terminal spatula 
with orifices (fig. 18, Gen). 


DISTRIBUTION: West Africa, Cameroon. 


REMARKS: This interesting species was found to be abundant in agricultural areas of 
the Sierra de las Minas and the Caribbean coast at Livinston. They live in the leaf 
litter or in rotten truncs of Cocos palms. B. kohleri was first discovered in Cameroon, 
a remarkable location for a member of this genus; since all the other species described 
are found in East Africa, South Asia and on the islands of the Wallacea. It is possible 
that this species was also introduced to Cameroon, all the localities where the species 
was found are close to the coast or to human settlements (Schmalfuss & Ferrara, 
1978, 1983 and 1985). Thus, the natural occurence of this species may still be 
unknown. On the other hand, Burmoniscus probably is paraphyletic and B. kohleri 
might not belong to this genus. B. kohleri is quite distinct from its congeners due to its 
minute size of maximally 3 mm and the number of spines along the medial margin of 
carpus | is reduced to two as opposed to three to four spines in other species of 
Burmoniscus. 

If this species is native to Cameroon, it is quite isolated from the African 
Oniscidean fauna by the position of the noduli laterales and the simple molar penicil, 
with one exception: Taiti & Ferrara (1980 and 1982) listed for the West African area a 


308 ANDREAS LEISTIKOW 


Fic. 16: Burmoniscus kohleri (Schmalfuss & Ferrara, 1978), d 2.7mm body length. Mdl/r left 
and right mandible, with detail of left pars intermedia; Mxp maxilliped, with detail of endite in 
rostral view; Mx1 maxillula, with detail of apical lateral endite in rostral view; Mx2 maxilla. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 309 


Fic. 17: Burmoniscus kohleri (Schmalfuss & Ferrara, 1978), & 2.7mm body length. Dac 
dactylus of pereopod 7 in rostral view; PE1-7 pereopods | to 7 in caudal or rostral (PE1) view; 
Sc] ornamental, longest and smallest sensory spines of carpus 1; Sp] medioproximal sensory 
spine of propus 1. 


310 ANDREAS LEISTIKOW 


N 


| 
| 
| 
| 
\ 
| 
| 
| 
| 


Fic. 18: Burmoniscus kohleri (Schmalfuss & Ferrara, 1978), d 2.7mm body length. Gen genital 
papilla; PL1-5 pleopods 1 to 5 in rostral view, with detail of endopodite 1 in caudal view. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 311 


genus which shares these characters. Zebrascia Verhoeff, 1942 was described on the 
basis of a single female of 3 mm body length from Fernando Po (Verhoeff, 1942). 
This genus now comprises three species; two of them from Cameroon (Schmalfuss & 
Ferrara, 1978). Interestingly, the pereopod 1 carpus is of similar shape in both 
Zebrascia and B. kohleri, and even the number and position of the sensory spines 
correspond in both taxa. The position of the noduli laterales with respect to the 
posterior margin of the coxal plates is similar in Zebrascia and B. kohleri (Schmalfuss 
& Ferrara, 1978, Taiti & Ferrara, 1986b). It may be well possible that B. kohleri 
belongs to a taxon close to Zebrascia; with the following synapomorphies: 


M Antenna-grooming brush covering half the medial margin of pereopod 
carpus | [antenna-grooming brush covering less than a third of carpus] 

@ Sensory spines of carpus | arranged in a 2-spine-pattern [sensory spines 
arranged in a 3-spine pattern] 


In the Indopacific species of Burmoniscus differ from B. kohleri by the 
pereopod | carpus being similar to the relations found in Androdeloscia (three-spine- 
pattern, see below and fig. 20, PEI) and the antenna-grooming brush being not so 
broad. To clarify this puzzling taxonomic situation, all species of Burmoniscus and 
Zebrascia have to be re-examined. 


Androdeloscia Leistikow, 1999 


REMARK: The genus Androdeloscia was recently instituted for the small members of 
the genus Prosekia Vandel, 1968; many species are found in northern South America 
(Leistikow, 1999), but there are no records from Central America until now. How- 
ever, two Central American species do belong to this genus, one originally described 
as a member of Philoscia (Mulaik, 1960), the other a new species from Guatemala. 
Beside /schioscia, Androdeloscia is the only genus of Crinocheta of philosciid appea- 
rance known to occur in Central America and northern South America (Leistikow, 
1997). 


Androdeloscia formosa (Mulaik, 1960) Figs 19-22 


Synonym: Philoscia formosa Mulaik, 1960 


Material examined: 1 d (body length 3 mm); 1 9: Guatemala, Petén, El Remate, 
Garden of the Casa de Don David, under coconuts and rotten leaves of Musa sp. and different 
species of deciduous trees, close to pasture at Lake Petén Itza, leg. 27.X.1998, A. Leistikow, 
author's collection; 3 4, several 2: Guatemala, Petén, causeway between Flores and Sta. Elena, 
100 m off Ciudad de Flores; in rotten Eichhornia sp. directly at lake shore, leg. 30.X.1998, A. 
Leistikow; author's collection and MNHG; 5 6, 4 ovigerous 9 : Guatemala, Petén, Sta. Elena, 
road to Grutas Actin Can; 250 m from entrance, under bark of cut trees, leg. 30.X.1998; A. 
Leistikow, author's collection; 3 d, 5 ® (Paratypes): Mexico, Tabasco, Palmillas, leg. 
18.VIII.1945, F. Bonet, IPNM 1102-E. 


Colour: Dorsally purplish brown with pale spots on pereon; medial line dark 
brown; white central stripe on pereonites I-IV; double line on pereonites V-VII; pleon 
unmarked. 


312 ANDREAS LEISTIKOW 


Cephalothorax: Linea frontalis and lamina frontalis lacking; linea supra- 
antennalis present; lateral lobes small; compound eyes consisting of eight ommatidia 
(fig. 19, Ctf). 

Pereon: Tegument smooth and shiny; coxal plates without gland pores; sulcus 
marginalis and noduli laterales present; the latter long and flagelliform; most dorsally 
inserted on coxal plate IV (fig. 19, Cx3/Cxp). 

Pleon: Narrower than pereon; neopleurae of pleonites 3 to 5 small; pleotelson 
with straight distal margin; bearing some tricorn-like setae. 

Appendages: 

Antennula: Three-articulate with prominent proximal article; distal joint 
bulbous; bearing two distinct sets of aesthetascs separated by a shallow depression 
(fig. 19, Anl). 

Antenna: Antennal peduncle composed of five articles with length ratio 1: 2: 2: 
3: 4; densely covered with tricorn-like setae; flagellum composed of three articles; 
distal one bearing prominent apical organ; as long as flagellar articles 1 and 2 together 
(fig. 19, An2). 

Mouth parts similar to following species. 

Pereopods: Pereopods slender; with setal tuft latero-distally on carpus; carpus 
1 with antennal-grooming brush; ornamental sensory spine serrate (fig. 20, Scl); 
dactylus with short inner claw and simple dactylar seta; interungual seta straight (fig. 
20, Dac). Sexual dimorphism: Male pereopod 7 merus with two small lobes medio- 
distally; prominent lobe below distal sensory spine, directed distally; covered with 
small scales (fig. 20, PE1-4; plate 21, PES-7). 

Pleopods: Exopodites rhomboid with two sensory spines laterally; endopodites 
subrectangular (fig. 22, PL1-5). Sexual dimorphism: Male pleopod 1 exopodite 
rounded with slight concavity laterally; endopodite rather bulky; basal part containing 
muscle M49 being half as broad as protopodite; laterally sinuous; distally with 
prominent hump; apex slightly bent laterally; with short medio-caudal row of spines 


groove for directing pleopod 2 exopodite; apex not elongate (fig. 22, PLS). 
Uropod: As in other species of the genus. 
Genital papilla: Rather short and slender compared to congeners (fig. 23, Gen). 


REMARKS: This species belongs to a distinct subtaxon of the genus Androdeloscia 
which is characterised by a sexual dimorphism of the male pereopod 7 merus. It bears 
several hooks on its medial margin similar to A. silvatica (Lemos de Castro & Souza, 
1986) and A. pseudosilvatica Leistikow, 1999 from the Caribbean region. Particularly 
A. pseudosilvatica resembles A. formosa: the male pereopod 7 merus has three hooks 
and the male pleopod | endopodite is bulbous at its base and falciform in the distal 
part. Both species are separated by several characters. In A. formosa the maxillula has 
a simple tooth more laterally, in A. pseudosilvatica the simple tooth is placed 
medially, the male pereopod 7 merus bears two sensory spines in vicinity of the 
proximal hook, in A. pseudosilvatica there is only one sensory spine, and the male 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 


313 


200um 


200m 
Cxp 
Fic. 19: Androdeloscia formosa (Mulaik, 1960), d 4mm body length. Anl antennula; An2 


antenna, with detail of apical organ; Ctf cephalothorax in frontal view; Cxp position of noduli 


laterales on coxal plates; Cx3 coxal plates3; Had habitus in dorsal view; Hal habitus in lateral 
view; Tel pleotelson. 


314 ANDREAS LEISTIKOW 


Fic. 20: Androdeloscia formosa (Mulaik, 1960), ¢ 4mm body length. Dac dactylus 1 in rostral 
view; PEI-4 pereopods 1-4 (caudal view), with detail of carpus 1 in rostral view; Scl orna- 
mental sensory spine of carpus 1; Sc2 longest sensory spine of carpus 2; Spl distal sensory 
spine of propus 1. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 315 


Fic. 21: Androdeloscia formosa (Mulaik, 1960), ¢ 4mm body length. PES-7 pereopods 5-7 in 
caudal view. 


316 ANDREAS LEISTIKOW 


PL2 PL3 


Fic. 22: Androdeloscia formosa (Mulaik, 1960), 4 4mm body length. Gen genital papilla; PL1- 
5 pleopods 1-5, rostral view, with detail of endopodite 1 in caudal view. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA Se 


pleopod 1 endopodite has the apex smooth and a lateral hump on half-length in A. 
formosa, wheras in A. pseudosilvatica, the apex bears some lateral hooks and there is 
no lateral hump. Additionally, the dactylar seta is acute in A. pseudosilvatica whereas 
the apex of the dactylar seta in A. formosa is pointed. A. pseudosilvatica might be the 
the adelphotaxon of A. formosa and A. silvatica, the latter share the lateral hump of 
the male pleopod | endopodite as a synapomorphy. 


Androdeloscia valdezi sp. n. Figs 23-26 


Material examined: & (holotype, body length 3 mm): Guatemala, Petén, causeway 
between Flores and Sta. Elena, 100 m off Ciudad de Flores, in rotten Eichhornia sp. at lake 
shore, leg. 30.X.1998, A. Leistikow, MNHG uncatalogised; Paratypes: 3 4; several 2: same 
data as for holotype, Guatemala, Petén, Sta. Elena, hill facing entrance to Grutas Actün Can, 
scrubs, loamy soil sparsely covered with leaf litter, leg. 30.X.1998, A. Leistikow, author's 
collection and UVG; 3 d, several 2: Guatemala; Zacapa, Rio Hondo, north of bridge over Rio 
Hondo, right bank, in leaf litter, leg. 18.X.1998; A. Leistikow, SMNS. 

DrAGNosis: Similar to most of its congeners, which do not show sexual dimorphism in 
the pereopods; differs in the shape of male pleopod | endopodite, which is pointed 
with lateral margin bearing several teeth. 

Colour: Mostly as in other species of the genus; dorsally reddish brown with 
light markings of muscle insertions; cephalothorax heavily spotted. 

Cephalothorax: Linea frontalis missing; lamina frontalis weak; linea supra- 
antennalis prominent; only slightly bent between antennal sockets; lateral lobes small; 
compound eyes consisting of about eight ommatidia (fig. 23, Ctf). 

Pereon: Tegument shiny; dorsum bearing scattered tricorn-like setae; coxal 
plates with sulcus marginalis and flagelliform noduli laterales; the latter more dorsally 
inserted on coxal plate IV (fig. 23, Cx3/Cxp). 

Pleon: Retracted from pereon; neopleurae attached; pleotelson with straight 
lateral margin; pointed; bearing short tricorn-like setae. 

Appendages: 

Antennula: Distal article apically with small tip and two aesthetascs; medially 
with tuft of about nine aesthetascs; medial article distinctly shorter than proximal one 
(fig. 23, Anl). 

Antenna: More slender than in preceding species; especially flagellum; distal 
article longest; proximal articles subequal in length; apical organ slender, longer than 
distal article (fig. 23, An2). 

Mandible: Molar penicil consisting of a three branches; pars intermedia 
bearing two penicils on left and one on right mandible; intermedial penicil slender 
(fig. 24, Mdl/r). 

Maxillula: Medial endite similar to preceding species; lateral endite with 4+6 
teeth; five of inner set cleft (fig. 24, Mx1). 

Maxilla: Lateral lobe only slightly broader than medial lobe; bearing pectinate 
scales; medial lobe sparsely covered with trichia; apically cuspidate (fig. 24, Mx2). 

Maxilliped: As in other species of Androdeloscia; basipodite in examined 
specimen broken (fig. 24, Mxp). 


318 ANDREAS LEISTIKOW 


Fic. 23: Androdeloscia valdezi sp. n., holotype & 3.5mm body length. Anl antennula; An2 
antenna, with detail of apical organ; Ctf cephalothorax in frontal view; Cxp position of noduli 
laterales on coxal plates; Cx3 coxal plate III; Had habitus in dorsal view; Hal habitus in lateral 
view; Tel pleotelson. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 319 


Fic. 24: Androdeloscia valdezi sp.n., holotype d 3.5mm body length. Mdl/r left and right 
mandible, with detail of pars intermedia; Mxp maxilliped, with detail of endite in rostral view; 
Mx1 maxillula, with detail of apex of lateral endite; Mx2 maxilla. 


320 ANDREAS LEISTIKOW 


Fic. 25: Androdeloscia valdezi sp. n., holotype & 3.5mm. Dac dactylus 1 in rostral view: 
PEI-7 pereopods 1 to 7 (caudal view), with details of carpus 1 in rostral and ischium 6 in 
caudal view; Scl ornamental sensory spine of carpus 1: Sp! distal sensory spine of propus 1. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 32] 


FIG. 26: Androdeloscia valdezi sp. n., holotype 4 3.5mm body length. Gen genital papilla; 
PLI-5 pleopods 1-5, rostral view, with details of endopodite 1 in caudal and rostral view. 


322 ANDREAS LEISTIKOW 


Pereopods: Rather slender; especially pereopod 1; carpus with antenna-groo- 
ming brush and serrate ornamental sensory spine (fig. 25, Sc1); pereopod 2 merus 
with hyaline fringe medially; dactylus with long inner claw and simple dactylar seta 
(fig. 25, Dac). Sexual dimorphism not evident (fig. 25, PE1-7). 

Pleopods: Pleopod exopodites 3 and 4 elongate rhomboid with two sensory 
spines laterally; exopodite 5 more or less triangular; endopodites bilobate; no respira- 
tory areas discernible in light microscope (fig. 26, PL1-5). Sexual dimorphism: Male 
pleopod 1 exopodite rounded; endopodite pointed with lateral margin bearing several 
teeth; basal part with muscle M49 less than half the length of endopodite; short caudal 
row of sensory spines (fig. 26, PL1). Pleopod 2 and 5 similar to preceding species 
(He227APE2/5); 

Uropod: As in other species of the genus. 

Genital papilla: Similar to preceding species (fig. 26, Gen). 

REMARKS: The new species is close to the “species-group D” of the genus Andro- 
deloscia with the autapomorphies: hyaline lobes on the male pleopod 1 endopodite 
reduced; maxillipedal basipodite slender, apically subrectangular (Leistikow, 1999). 
The exact systematic position within this group is difficult to access: the male pleopod 
1 endopodite in A. valdezi is similar to that of A. conipus Leistikow, 1999 and related 
species, but this similarity is possibly a symplesiomorphy. A. poeppigi Leistikow, 
1999 and A. malleus Leistikow, 1999 form the sister group of A. conipus and related 
species. The former have the more derived endopodites | within species-group D. The 
caudo-medial row of spines is reduced and there are two subapical lobes, synapo- 
morphic characters of A. poeppigi and A. malleus. For the time being, the systematic 
positon of the new species remains open to debate. 


ACKNOWLEDGEMENTS 


The author wishes to thank the following persons: Prof. Dr J.W. Wägele, 
Ruhr-Universität Bochum, for critically discussing the results of this work; Dr M. 
Dix, Mrs Mayra Maldonado and Mr José Rodolfo Valdez Barilla, Universidad del 
Valle de Guatemala and Mrs. Beate Fasselt for their support during the field trip in 
Guatemala. For financial support of the field trip, the author is indebted to the 
Studienstiftung des Deutschen Volkes. This study is dedicated to the victims of the 
Hurricane “Mitch”, which devastated the northern Central American region in 
October 1998. 


REFERENCES 


ARAUIO, P. B., Buckup, L. AND BOND-BUCKUP, C. 1996. Isopodos terrestres (Crustacea, 
Oniscidea) de Santa Catarina e Rio Grande do Sul. /heringia Serie Zoologia 81: 111- 
138. 

BRIAN, A. 1929. Descrizione di un nuovo genere di isopode terrestre troglobio raccolto dal 
Prof. Silvestri. Bollettino del Laboratorio de Zoologia generale ed Agraria della 
Facclta Agraria in Portici 22: 188-197. 


TERRESTRIAL ISOPODA FROM MEXICO AND GUATEMALA 323 


ERHARD, F. 1997. Das pleonale Skelet-Muskel-System von Titanethes albus (Synocheta) und 
weiterer Taxa der Oniscidea (Isopoda) mit Schlußfolgerungen zur Phylogenie der 
Landasseln. Stuttgarter Beiträge zur Naturkunde Serie A (Biologie) 550: 1-70. 

FERRARA, F. & SCHMALFUSS, H. 1983. New isopod material from Southwest Cameroon, with 
descriptions of 13 new species. Stuttgarter Beiträge zur Naturkunde. Serie A (Biologie) 
360: 1-43. 

FERRARA, F. & SCHMALFUSS, H. 1985. Terestrial isopods from West Africa. Part 4: Addenda 
and conclusions. Monitore zoologico italiano (n. s.) suppl. 20: 55-120. 

FERRARA, F. & TAITI, S. 1981. Isopodi terrestri delle isole Andamane. Memorie del Museo 
Civico di Storia Naturale in Verona 8: 459-492. 

FERRARA, F. & TAITI, S. 1986. The validity of Anchiphiloscia Stebbing, 1908 (Crustacea Iso- 
poda, Oniscidea). Monitore zoologico italiano (new series) suppl. 21: 149-167. 

LEISTIKOW, A. 1997. Terrestrial Isopods from Costa Rica and redescription of /schioscia 
variegata (Dollfus, 1893). Canadian Journal of Zoology 75: 1415-1464. 

LEISTIKOW, A. 1999. Androdeloscia gen. n., a new genus of South American terrestrial isopods 
with description of 13 new species (Crustacea: Isopoda: “Philosciidae”). Revue suisse 
de Zoologie 106: 1-92. 

Miers, E. J. 1877. On a collection of Crustacea, Decapoda and Isopoda, chiefly from South 
America. Proceedings of the zoological Society of London 1877: 653-679. 

Mu alk, S. 1960. Contribuciön al conocimiento de los isopodos terrestres de México. Revista 
de la Sociedad de México de Historia Natural 21: 79-292. 

RICHARDSON, H. 1907. A new terrestrial isopod from Guatemala, type of a new genus. 
Proceedings of the U.S. National Museum 32: 447-450. 

RICHARDSON, H. 1910. Description of a new terrestrial isopod from Guatemala. Proceedings of 
the U.S. National Museum 37: 495-497. 

SCHMALFUSS, H. 1989. Phylogenetics in Oniscidea. Monitore zoologico italiano (n.s.) 4: 3-27. 

SCHMALFUSS, H. & FERRARA, F. 1978. Terrestrial isopods from West Africa. Part 2: Families 
Tylidae, Ligiidae, Trichoniscidae, Styloniscidae, Rhyscotidae, Halophilosciidae, Philo- 
sciidae, Platyarthridae, Trachelipidae, Porcellionidae, Armadillidiidae. Monitore zoolo- 
gico italiano (n. s.) suppl. 11(2): 15-97. 

SCHULTZ, G. A. 1977. Two blind species, one new, of terrestrial isopod crustaceans (Onis- 
coidea) from Yucat-n and Guatemala. Association of Mexican Cave Studies Bulletin 6: 
9-13. 

STROUHAL, H. 1966. Eine neue halophile Stenophiloscia aus dem Rotmeergebiete. Annalen des 
Naturhistorischen Museums in Wien 69: 323-333. 

TAITI, S. & FERRARA, F. 1980. The family Philosciidae in Africa, south of the Sahara. Monitore 
zoologico italiano (n. s.) 13: 53-98. 

TAITI, S. & FERRARA, F. 1982. Revision of the family Philosciidae from South Africa. Annals of 
the South African Museum 90: 1-48. 

TAITI, S. & FERRARA, F. 1986a. Taxonomic revision of the genus Littorophiloscia Hatch, 1947 
(Crustacea, Isopoda, Oniscidea) with description of six new species. Journal of Natural 
History 20: 1347-1380. 

TAITI, S. & FERRARA, F. 1986b. Terrestrial Isopoda from the oriental region. I. The genus 
Burmoniscus Collinge, 1914 (Philosciidae). Monitore zoologico italiano (n. s.) suppl. 
2125-133: 

VANDEL, A. 1973a. Les isopodes terrestres (Oniscoidea) de la Mélanésie. Zoologische verhand- 
lingen 125: 1-160. 

VANDEL, A. 1973b. Les isopodes terrestres de |’ Australie, étude systématique et biogéo- 
graphique. Mémoires du Museum National d’Histoire Naturelle, n. s., A (Zoologie) 
SALI 

VERHOEFF, K. W. 1942. Landisopoden von Fernando Po. Zoologischer Anzeiger 137: 84-98. 


REVUE SUISSE DE ZOOLOGIE 107 (2): 325-334; juin 2000 


Euphalerus clitoriae sp. n., a new psyllid species from 
Clitoria fairchildiana (Fabaceae, Papilionoideae), and notes 
on other Euphalerus spp. (Hemiptera, Psylloidea) 


Daniel BURCKHARDT! & Mariängela GUAJARA? 

| Naturhistorisches Museum, Augustinergasse 2, CH-4001 Basel, Switzerland. 
E-mail: daniel.burckhardt@unibas.ch 

2 Universidade Federal Rural do Rio de Janeiro, CPG em Fitotecnia 
Seropédica, RJ — 23.851-970, Brazil. 
E-mail: guajara@ufrrj.br 


Euphalerus clitoriae sp. n., a new psyllid species from Clitoria fair- 
childiana (Fabaceae, Papilionoideae), and notes on other Euphalerus 
spp. (Hemiptera, Psylloidea). - Euphalerus clitoriae sp. n. from Brazil, a 
species developing under waxy filamentous secretions on the leaves of 
Clitoria fairchildiana, ıs described and illustrated. Its biology is briefly 
outlined and compared to that of its closest relatives, viz. E. nidicola from 
Peru and E. maya from Belize. Differences to these two species are 
discussed. Based on the study of types, the genitalia of E. nidicola, E. 
antillensis and E. ostreoides are illustrated completing their insufficient 
original descriptions. 


Key-words: Psylloidea - Fabaceae - Brazil - taxonomy - biology. 


INTRODUCTION 


The hemipterous jumping plant-lice or psylloids are usually highly host specific 
and thus a potentially suitable taxon for coevolutionary studies on insect — plant 
relationships (Burckhardt & Basset, 2000). The necessary taxonomic and phylogenetic 
base is, however, often insufficient or completely lacking. The genus Euphalerus is a 
good example. It was erected for the Carribean species E. nidifex whose larvae form 
lerps or “nests”, i.e. waxy coverings, on Fish poison Bark or Jamaican dogwood 
(Piscidia piscipula and Piscidia carthegenensis, Fabaceae, Papilionoideae, Tephro- 
sieae) (Schwarz, 1904; Russell, 1971). Subsequently, a series of mostly tropical New 
and Old World species were referred to the genus making it extremely artificial. 
According to Hollis & Martin (1997), the New World species forming lerps or galls on 
the leaves of Fabaceae make up true Euphalerus (Psyllidae, Euphalerinae). The other 
New World species, associated with Ceanothus (Rhamnaceae) and Cerocarpus (Rosa- 
ceae), are unrelated and belong to the Arytaininae/Psyllinae (Psyllidae) complex. The 
Old World species, finally, are probably referable to the euphalerine genera Colo- 
phorina Capener and Euryconus Aulmann. 


Manuscript accepted 08.11.1999 


326 DANIEL BURCKHARDT & MARIANGELA GUAJARA 


Twenty two species are currently known in Euphalerus s. str. The vast majority 
is associated with Lonchocarpus (Papilionoideae, Tephrosieae). One species is each on 
Pithecellobium (Mimosoideae, Ingeae), Erythrina (Papilionoideae, Phaseoleae) and 
Piscidia (Papilionoideae, Tephrosieae) respectively, two species have unknown and one 
doubtful (Karwinskia, Rhamnaceae) host records. Nineteen species are recorded from 
Central America with one species extending to Florida, and only three are also or 
exclusively known from South America (Brazil 2, Peru 1) (Hollis & Martin, 1997). 

The present paper describes a new species from Brazil developing on Clitoria 
fairchildiana (Papilionoideae, Phaseoleae). The new species broadens the known host 
range of Euphalerus, and suggests that the genus may be more diverse in tropical South 
America than currently estimated. It is closely related to the gall-forming E. nidicola 
Tuthill from Peru whose larvae are covered in waxy secretions, and the lerp-inhabiting 
E. maya Hollis & Martin from Belize. Hollis & Martin (1997) provided detailed 
descriptions of the species attacking Lochocarpus in Belize. Here we give illustrations 
of the male and female genitalia of E. antillensis Caldwell & Martorell and E. nidicola 
Tuthill, and of the male genitalia of E. ostreoides Crawford whose original descriptions 
lack sufficient detail and which were not treated by Hollis & Martin (1997). 


MATERIAL AND METHODS 


The morphological terminology follows Hollis (1976) and Hollis & Martin 
(1997). Measurements and some drawings were made from slide mounted material. The 
drawings for E. antillensis and E. nidicola were made from temporary mounts in 
glycerine. 

Material is cited from following institutions: Naturhistorisches Museum Basel, 
Switzerland - NHMB; Angelo Moreira da Costa Lima Entomological Collection, Sero- 
pédica, Brazil - CLEC; Natural History Museum, London, UK - BMNH; United States 
National Museum of Natural History, Washington, DC, USA (psylloid collection in 
USDA, Beltsville, MD) - USNM; Muséum d’histoire naturelle, Geneva, Switzerland - 
MHNG. 


TAXONOMIC TREATMENT 
Euphalerus clitoriae sp. n. Figs 1-3, 13, 16-23 


Holotype & , Brazil: State of Rio de Janeiro, Campus and residential area of the Univer- 
sidade Federal Rural do Rio de Janeiro, Seropédica, 22°44' S 43°43' W, 9.1x.1999, Clitoria 
fairchildiana (M. Guajara), dry mounted (NHMB). 

Paratypes, Brazil: 20 d, 20 9, 10 larvae or larval skins, same data as holotype, dry 
mounted (NHMB, CLEC, BMNH, USNM, MHNG); 6 d, 6 ©, 4 larvae, same data but 30.v. 
1999, dry and slide mounted (NHMB, CLEC, BMNH). 


Material not included in type series. Brazil, many adults and some larvae, same data as 
holotype but 30.v.1999 and 9.1x.1999, preserved in 70 % alcohol (NHMB). 

Adult. Body dirty greyish white with small blackish dots, in mature specimens 
thorax dorsally, abdomen and genitalia dark. Antennal segments 1-8 light with dark 
apices, segments 9 and 10 entirely dark. Forewing membrane transparent, colourless, 
with blackish dots as in fig. 16, middle of cells very weakly infuscate; veins whitish 
with black dots. Legs whitish with dark dots, tarsi blackish. 


EUPHALERUS CLITORIAE 327 


Small species (see measurements below). Genal processes longer than vertex 
along mid-line, conical, subacute apically (fig. 17). Antenna 1.97-2.13 times as long as 
head width, segment 3 (1.32 times) longer than segment 8. Ultimate two rostral 
segments 0.41-0.43 times as long as head width. Forewing (fig. 16) 2.24-2.31 times as 
long as wide, 3.67-3.93 times as long as head width; pterostigma moderately long, 0.5 
times as long as vein Rs; surface spinules leaving broad spinule-free stripes along the 
veins; reduced in basal half of cell r, and r,, occupying only a small area in cell c+sc. 
Metatibia 0.93-0.97 times as long as head width, bearing a small basal spine and an 
incomplete crown of apical spurs. 

Male proctiger (fig. 1) bulbous, lacking lateral lobes. Paramere in profile (fig. 2) 
with each an anterior and a posterior lobe. Distal portion of aedeagus (fig. 3) hook- 
shaped; apex of terminal tube of ductus ejaculatorius truncate. 

Female proctiger (fig. 13) more or less evenly tapering, dorsal margin weakly 
concave, apex narrowly rounded; 1.00 times as long as head width, 3.21 times as long 
as circumanal ring, 1.56 times as long as subgenital plate. 

Measurements in mm (1 6, 1 9). Head width 0.63; antenna length 1.25-1.35; 
forewing length 2.33-2.50; male proctiger length 0.26; paramere length 0.23; length of 
distal segment of aedeagus 0.23; female proctiger length 0.63. 

Fifth instar larva (fig. 18). Antenna 8-segmented with one rhinarium on each of 
segments 3, 5, 7 and 8. Tibiatarsus of foreleg with one large apical and one smaller 
subapical spur (fig. 19); tibiotarsi of mid and hindlegs with one large and two small 
spurs. Subgenital plate with two apical processes bearing three teeth each (fig. 20). 

Host plant. Clitoria fairchildiana Howard (= racemosa) (Fabaceae). 

Biology. The eggs are laid along the veins of the leaves. The larvae sit on the 
leaves and stems hidden under white coverings consisting of waxy hair-like filaments 
(fig. 21). There is no sign of deformations on the leaves (fig. 22). When the larval 
density is high, the leaves turn yellow and drop. Strongly infested trees can loose all 
their leaves (fig. 23). 

Comments. E. clitoriae, which is a member of the E. ostreoides species group as 
defined by Hollis & Martin (1997), is closely related to E. nidicola Tuthill, 1959 from 
Peru, and E. maya Hollis & Martin, 1987 from Belize based on following characters. 
Small sized Euphalerus spp.; head with long, conical genal processes; antenna longer 
than 1.5 times head width; forewing clear, bearing dark spots lacking transverse bands; 
metatibia with a small basal spine and an incomplete crown of apical spurs; male 
proctiger bulbous without lateral lobes; paramere complex, irregularly S-shaped with a 
group of long thick setae in the middle of the foremargin and a group of heavily 
sclerotised peg setae on the inner surface of the posterior lobe; female subgenital plate, 
in profile, with angular ventral margin. 

E. clitoriae differs from E. maya in the slightly larger body dimensions, the 
longer antenna, the shorter pterostigma of the forewing, the details in the genital 
structure and the apex of the larval caudal plate which has three points on each tubercle 
instead of two. E. clitoriae is differentiated from E. nidicola by the much more 
expanded, almost black body coloration, which is orange to light brownish in E. 
nidicola, the blackish dots on the forewings which are brown in E. nidicola, the shorter 


328 DANIEL BURCKHARDT & MARIANGELA GUAJARA 


Fics 1-6 


Euphalerus spp. 1, 4: Male genitalia, in profile; 2, 5: paramere, inner face; 3, 6: distal portion of 
aedeagus. 1-3: E. clitoriae sp. n.; 4-6: E. nidicola Tuthill. Scale lines: figs 1, 4 = 0.2 mm, figs 2, 
3, 4,6=0.1 mm. 


EUPHALERUS CLITORIAE 329 


0.2 mm 


0.3 mm 


Fics 7-12 


Euphalerus spp. 7, 10: Male genitalia, in profile; 8, 11: paramere, inner face; 9, 12: distal portion 
of aedeagus. 7-9: E. antillensis Caldwell & Martorell; 10-12: E. ostreoides Crawford. Scale lines: 
fig. 7 = 0.2 mm, figs 8, 9 = 0.1 mm, fig. 10 = 0.3 mm, figs 11, 12 = 0.2 mm. 


antenna, the more reduced fields of surface spinules in the forewing, the details of the 
genitalia (for E. nidicola see figs 4-6, 14); the female proctiger is slightly shorter and 
apically more thickened in E. clitoriae but longer and apically pointed in E. nidicola. 
The three species differ also in their geographical and host plant ranges. The 
larval biology of E. clitoriae is intermediate. It shares with E. nidicola waxy filamen- 


330 DANIEL BURCKHARDT & MARIANGELA GUAJARA 


tous coverings but differs by not causing depressions on the leaves. This character is as 
in E. maya which is characterised by the production of a lerp. 


Euphalerus antillensis Caldwell & Martorell Figs 7-9, 15 


Euphalerus antillensis Caldwell & Martorell, 1951: 612. Holotype d, Puerto Rico: Guänica, on 
the Guanica - Guänica Central Road, in front of stone quarry, 28.viii.1947, breeding abundantly 
on the undersides of the leaves of “geno-geno”, Lonchocarpus domingensis (Caldwell & 
Martorell) (USNM). 


Material examined. Puerto Rico: 1 d, 1 2 paratypes, same data as holotype (USNM). 


Adult. Member of the nidifex species group (as defined by Hollis & Martin, 
1997). Metatibia with a small basal spine and grouped apical spurs. 

Male proctiger (fig. 7) with narrow lateral lobes. Paramere in profile (fig. 8) 
lamellar, obliquely truncate apically, inner surface covered in long setae. Distal portion 
of aedeagus (fig. 9) with oval apical dilatation, rounded apically; apex of terminal tube 
of ductus ejaculatorius rounded. 

Female proctiger (fig. 15) strongly tapering to the middle, digitiform in apical 
half, dorsal margin strongly concave, apex narrowly rounded. 


Euphalerus nidicola Tuthill Figs 4-6, 14 
Euphalerus nidicola Tuthill, 1959: 6. Holotype d, Peru: few km below La Merced, 1.1.1947, on 
“oropel”, Erythrina sp. (Tuthill) (USNM). 

Material examined. Peru: holotype à , allotype 2,1 d,1 © paratypes (USNM). 


Adult. Member of the ostreoides species group (as defined by Hollis & Martin, 
1997). 

Male proctiger (fig. 4) bulbous, lacking lateral lobes. Paramere in profile (fig. 5) 
with each an anterior and a posterior lobe. Distal portion of aedeagus (fig. 6) hook- 
shaped; apex of terminal tube of ductus ejaculatorius truncate. 

Female proctiger (fig. 14) irregularly tapering, dorsal margin weakly but 
distinctly concave, apex subacute. 

Fifth instar larva. Tuthill (1959) mentioned the presence of many larvae. In the 
collection of the USNM we could, however, find only the adult types. 


Euphalerus ostreoides Crawford Figs 10-12 


Psyllid, Tavares, 1920: 124; 1922: plate 19 figs 3-5. Galls on the leaves of an undetermined 
species of Leguminosae (Timb6), Brazil: Rio de Janeiro State, Nova Friburgo; Rio de Janeiro 
State, surroundings of Rio de Janeiro; Sao Paulo State, Itu; between Rio Vermelho and the city of 
Bahia. 

Euphalerus ostreoides Crawford, 1925: 62. Syntypes 2 d, 1 ®, Brazil: Sao Paulo, Itu, making 
very peculiar galls on the leaves of an undetermined species of Leguminosae (Tavares) (USNM). 
Costa Lima, 1942: 107; Silva et al., 1968: 201; Russell, 1971: 10; Hollis & Martin, 1997: 241. 


Material examined. Brazil: syntype d, Sao Paulo, Itu, (Tavares) (slide mounted, USNM). 


Adult. Member of the ostreoides species group (as defined by Hollis & Martin, 
1997). Large species (see measurements below). Genal processes shorter than vertex 
along mid-line, broadly rounded apically. Antenna 2.32 times as long as head width, 
segment 3 about as long as segment 8. Ultimate two rostral segments 0.39 times as long 


EUPHALERUS CLITORIAE 331 


= 
Ve 4, 
0.2 mm a > 
“2 7; "pt 
Ni i fe 


Fics 13-15 


Euphalerus spp., female genitalia, in profile. 13: E. clitoriae sp. n.; 14: E. nidicola Tuthill; 15: 
E. antillensis Caldwell & Martorell. Scale line = 0.2 mm. 


332 DANIEL BURCKHARDT & MARIANGELA GUAJARA 


c 
E 
e 
ei 

a 


Fics 16-20 


Euphalerus clitoriae sp. n. 16: forewing; 17: head, dorsal view; 18: fifth instar larva, left dorsal, 
right ventral view; 19: apex of foreleg of fifth instar larva; 20: apex of caudal plate, dorsal view. 
Scale lines: fig. 16 = 1.0 mm, fig. 17 = 0.3 mm, fig. 18 = 0.3 mm, figs 19, 20 = 0.1 mm. 


333 


EUPHALERUS CLITORIAE 


‘u ‘ds 2011012 snaajoyd 
-n7 JO uonsajur AAvay 0) ANP soABa] OU Jsouufe 
YUM DUDIP]IY9AMS DIIOIMI JO IL ‘ET OM 


"SUOHEULIOFOP oxI]-11d Buryory 
S9AB9] DUDIP]1Y241Df D140], UO ‘u ‘ds 2014017179 
snıa]oydn7 JO BAIL] P919A09 XEM ‘77 ‘DIA 


‘SU9JS pue SOABO] DUDIP}1Y9410f 
D14011]9) UO "u ‘ds 20140179 Sh12]Dydn JO SVAIE] 
au} Aq SUOH2199$ aNIj-Iey ÂXEM NUM ‘IT DIA 


334 DANIEL BURCKHARDT & MARIANGELA GUAJARA 


as head width. Forewing 2.34 times as long as wide, 3.06 times as long as head width; 
pterostigma relatively short, 2.6 times as long as vein Rs; surface spinules present in 
apical part of all cells except for c+sc, leaving broad spinule-free stripes along the 
veins; radular spinules covering broad triangular patches along the wing margin in cells 
Ty, 1, Mj, m, and cu,. Metatibia 0.77 times as long as head width, bearing a conspi- 
cuous basal spine and an incomplete crown of apical spurs. 

Male proctiger (fig. 10) with distinct lateral lobes. Paramere in profile (fig. 11) 
with a large posterior lobe, inner face with an anterior field of longer, more spaced setae 
and a posterior field with peg-like, densely spaced setae. Distal portion of aedeagus 
(fig. 12) rounded apically, with inflated apical half; apex of terminal tube of ductus 
ejaculatorius truncate. 

Female unavailable. 

Measurements in mm (1 9). Head width 1.15; antenna length 2.68; forewing 
length 3.54; male proctiger length 0.47; paramere length 0.47; length of distal segment 
of aedeagus 0.47. 


ACKNOWLEDGEMENTS 


We thank D. Miller (USDA, Beltsville Md, USA) for the loan of material, D. 
Hollis (BMNH) for comments on Euphalerus and on a manuscript draft, and A. G. de 
Carvalho (Seropédica) for pointing out the new species and suggesting its study. 


REFERENCES 


BURCKHARDT, D. & BASSET, Y. 2000. The jumping plant-lice (Hemiptera, Psylloidea) associated 
with Schinus (Anacardiaceae): systematics, biogeography and host plant relationships. 
Journal of Natural History 34: 57-155. 

CALDWELL, J. S. & MARTORELL, L. F. 1951. A brief review of the Psylliidae of Puerto Rico 
(Homoptera). Annals of the Entomological Society of America 44: 603-613. 

Costa LIMA, A. DA. 1942. Homopteros. Superfamilia Psylloidea. Insetos do Brasil 3: 94-111. 

CRAWFORD, D. L. 1925. Psyllidae of South America. Broteria 12: 56-74. 

HoLLis, D. 1976. Jumping plant lice of the tribe Ciriacremini (Homoptera: Psylloidea) in the 
Ethiopian Region. Bulletin of the British Museum (Natural History) (Entomology) 43: 
1-83. 

Ho tis, D. & MARTIN, J. 1997. Jumping plantlice (Insecta: Hemiptera) attacking Lonchocarpus 
species (Leguminosae), including ‘Black Cabbage Bark’, in Belize. Journal of Natural 
History 31: 237-267. 

RUSSELL, L. M. 1971. Notes on Euphalerus nidifex Schwarz and related nest-making New World 
psyllids (Homoptera: Psyllidae). Florida Entomologist 54: 3-12. 

SCHWARZ, E. A. 1904. Notes on North American Psyllidae. Part I. 2. Description of the nest- 
constructing Psyllid. Proceedings of the entomological Society of Washington 6: 234-245. 

SILVA, A. G. D’ ARAUJO E, GONÇALES C. R., GALVAO D. M., GONGALES A. J. L., GOMES J., SILVA, 
M. DO N. & SIMONI, L. DE. 1968. Quarto catalogo dos insetos que vivem nas plantas do 
Brasil, seus parasitos e predadores. Parte II - I tomo, insetos, hospedeiros e inimigos 
naturais, Rio de Janeiro, xxiv + (iv) + 622 pp. 

TAVARES, J. S. 1920. Cecidologia Brazileira. Broteria, Série Zoologica 18: 82-125. 

TAVARES, J. S. 1922. Cecidologia Brazileira. Broteria, Série Zoologica 20: 5-48. 

TUTHILL, L. D. 1959. Los Psyllidae del Peru Central (Insecta: Homoptera). Revista Peruana de 
Entomologia Agricola 2: 1-27. 


REVUE SUISSE DE ZOOLOGIE 107 (2): 335-349; juin 2000 


Zur Biologie des “Eingeweidefisches” 
Carapus acus (Brünnich, 1768) (Carapidae, Teleostei), 
mit Hinweisen auf eine nicht-parasitische Ernährung! 


Kerstin KLOSS und Wolfgang PFEIFFER 
Zoologisches Institut der Universität Tübingen 
Auf der Morgenstelle 28, D-72076 Tübingen, 
und 

Laboratoire Arago, Universite Paris VI 

F - 66650 Banyuls-sur-mer. 


On the biology of the pearlfish Carapus acus (Brünnich, 1768) (Cara- 
pidae, Teleostei), with indications of a non-parasitic nutrition. - 450 
Stichopus regalis (Cuvier), caught with a trawl-net at a depth of 70-110 m 
off Banyuls-sur-mer (France) contained 38 Carapus acus. We found only 
two pearlfish in 85 Holothuria tubulosa Gmelin caught in less than 10 m 
of seawater. We kept pearlfish in aquaria and analysed their behaviour by 
means of computer analysis of video records. The manner by which the 
fish enter their hosts was studied (see summary). In the wild most spe- 
cimens of C. acus are found in S. regalis, yet in our aquaria C. acus only 
entered H. tubulosa if they could choose between both species. We studied 
the stomach contents of C. acus and found crustaceans. The stomachs 
examined did not contain any holothurian tissue. Our results give reason to 
agree with Emery (1880), who stated that C. acus is an inquiline, to whom 
the host holothurian serves as a shelter only. 


Key-words: pearlfish - Carapus acus - Carapidae - Teleostei - host holo- 
thurian - Stichopus regalis - Holothuria tubulosa - stomach contents - 
behaviour - parasitism - inquilinism. 


EINLEITUNG 


Der “Eingeweidefisch” Carapus acus (Briinnich, 1768) (gängiges Synonym 
Fierasfer acus) ist in jeder Beziehung außergewöhnlich und erregte aufgrund seiner 
besonderen epidermalen Kolbenzellen bereits 1957 die Aufmerksamkeit des Zweit- 
Autors (W. P.). Diese Zellen ähneln weitgehend den Schreckstoffzellen der Ostario- 
physi (Pfeiffer, 1960). Karl von Frisch, Doktorvater des Zweit-Autors, forderte diesen 
bei seiner Abreise an die Zoologische Station in Neapel Anfang 1957 auf, den 
Mageninhalt von C. acus zu untersuchen. Dies geschah an 2 von Pierre Tardent zur 


! In memoriam Prof. Dr. Pierre Tardent. 
Manuskript angenommen 09.11.1999 


336 KERSTIN KLOSS UND WOLFGANG PFEIFFER 


Verfügung gestellten Exemplaren; beide enthielten Reste von Kleinkrebsen, wie an den 
schwarzen Komplexaugen deutlich zu erkennen war. Dieser Befund wurde Alfred 
Kaestner mitgeteilt und ist in dessen Lehrbuch nachzulesen (Kaestner, 1963, S. 1235). 
Zwanzig Exkursionen an das Laboratoire Arago in Banyuls-sur-mer (1963 und 1980 - 
1999) ermöglichten C. acus zu beobachten, zu sammeln und mit ihm zu experimen- 
tieren. Im Mittelpunkt des Interesses standen dabei die Fragen nach seiner Häufigkeit 
und Verbreitung, seinem Habitat und der Bevorzugung verschiedener Seewalzen- 
Species, der Art und Weise seines Eindringens, der Verbleibdauer im Wirt und nicht 
zuletzt seiner Ernährung. Handelt es sich bei C. acus tatsächlich um einen Nahrungs- 
parasiten, der Teile der Seewalze frißt, wie von Arnold (1953) behauptet aber nicht 
bewiesen und von der Sekundarliteratur kritiklos übernommen worden ist? Oder ist C. 
acus ein Inquilinist, dem die Seewalze lediglich als Wohnhöhle dient, wie von Emery 
(1880) beschrieben? Diesen Fragen sind wir im folgenden nachgegangen. 


MATERIAL UND METHODEN 


Alle Exemplare von C. acus wurden an der Küste vor Banyuls-sur-mer (Frank- 
reich) gesammelt: 38 C. acus stammten aus 450 Stichopus regalis (Cuvier) (70 - 110 m 
Meerestiefe), nur 2 aus 85 Holothuria tubulosa Gmelin (weniger als 10 m Meerestiefe). 
Demgegenüber enthielten 142 Holothuria polii Delle Chiaje und 63 H. tubulosa aus der 
Bucht von Port de la Selva (weniger als 10 m Wassertiefe) keinen einzigen C. acus. 
Weil die Fische bisher fast ausschließlich in ihren Wirten und nahezu nie frei- 
schwimmend gefunden worden waren, konzentrierten wir uns beim Fang auf die als 
Wirte bekannten Aspidochirota S. regalis, H. tubulosa und H. polii. Die Seewalzen 
wurden mittels eines Schleppnetzes gesammelt, das in 70 - 110 m Tiefe 60 - 90 min 
lang parallel zur Tiefenlinie über den Grund gezogen wurde. Die dabei gefangenen 
S. regalis - die beiden anderen Species kommen in dieser Tiefe vor Banyuls-sur-mer 
nur äußerst selten vor - wurden an Bord in einem Behälter mit Meerwasserdurchlauf 
aufbewahrt und an Land aufgeschnitten, um zu prüfen ob sie C. acus enthielten. Beim 
Eröffnen ihrer physiologischen Unterseite mit einer kräftigen Präparierschere war es 
unerheblich, ob der Schnittansatz im Maul oder im After erfolgte. Die Fische können 
sich aufgrund ihrer flexiblen Wirbelsäule an dem der Einschnittstelle gegenüber- 
liegenden Ende zu einem kleinen Knäuel zusammenrollen, so daß sie leicht zu 
übersehen sind (Abb. 2e). Wegen des grobmaschigen Schleppnetzes war nicht gewähr- 
leistet, daß freischwimmende C. acus oder solche, die von ihrer Seewalze mitsamt den 
Eingeweiden ausgestoßen worden waren, mitgefangen wurden. Um das Entkommen 
der Fische zu vermeiden, wurden H. tubulosa und H. polii von Hand gesammelt und 
unter Wasser in Plastiksäcken aufbewahrt. Beim Aufschneiden war erhöhte Vorsicht 
geboten, da die Arten von Holothuria eine wesentlich stärkere Hautmuskulatur besitzen 
als S. regalis und sich damit zu einer extrem harten Kugel zusammenziehen können. 
Alle C. acus wurden zusammen mit Seewalzen in Meerwasseraquarien gehalten. Die 
Versuche wurden in einem Glasaquarium (40 x 20 x 25 cm) und in einem Plastik- 
aquarıum (50 x 30 x 30 cm) durchgeführt. Um es den wegen der Fangbedingungen 
durch Dekompressionsschäden geschwächten Fischen zu ermöglichen in die Seewalze 


BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 337 


zu gelangen, wurde der Wasserspiegel während der ersten Tage nach dem Fang bis auf 
Höhe des Afters der Seewalze gesenkt. Für die Beobachtung des Eindringens wurde 
jeweils ein C. acus zusammen mit einer H. tubulosa oder einem S. regalis in ein 
Aquarium gesetzt. Bei den Wahlversuchen bekam der Fisch immer eine H. tubulosa 
und einen S. regalis, die wahllos aus der Menge der vorhandenen Seewalzen heraus- 
gegriffen worden waren. Wenn C. acus in einer Seewalze verschwunden war, wurde 
das Verhalten des Wirtes mindestens 5 min weiter beobachtet, um zu prüfen ob der 
Fisch ihn wieder verläßt. Nach Versuchsende wurde C. acus mit der Seewalze wieder 
in das Hälterungsbecken zurückgesetzt. Die Dokumentation der Versuche erfolgte 
exemplarisch mittels Videokamera (Typ: Sony Video 8) und Photoapparat (Typ: Canon 
EOS 100). Ansonsten wurden schriftliche Protokolle erstellt. Um zu klären welche 
Faktoren für die Art des Einstiegs in die Seewalze verantwortlich sind, wurde das vor- 
und rückwärtige Eindringen der Fische in Einzelbildsequenzen aufgelöst. Zur 
Auswertung dieser Sequenzen dienten ein Videorecorder (Typ: Panasonic) und ein 
Computer (Programm: Scancam). 

Auch alle für die anatomischen und histologischen Untersuchungen verwen- 
deten C. acus Exemplare entstammten S. regalis aus 70 - 110 m Meerestiefe. Die 
Fische wurden in 10% Formalin oder nach Bouin fixiert und später in 70% Ethyl- 
alkohol überführt. Sie wurden anschließend vermessen, der Länge nach sortiert, 
numeriert und ihr Geschlecht anhand der Gonaden bestimmt. Im Labor wurden die 
Mägen von in Formalin fixierten C. acus unter dem Binokular geöffnet und auf ihren 
Inhalt untersucht. Die dabei gefundenen Kleinkrebse wurden mit Boraxkarmin gefärbt 
und in Kanadabalsam eingeschlossen. Bei nach Bouin fixierten Fischen wurden die 
Magen ganz herauspräpariert und in Paraffin eingebettet. Von den Paraffinblöcken 
wurden 7um dicke Schnitte angefertigt, mit Hämalaun-Eosin oder Azan gefärbt und 
unter einem Zeiss-Mikroskop ausgewertet und photographiert. 


ERGEBNISSE 


Die Häufigkeit von C. acus in S. regalis war in den verschiedenen Jahren sehr 
unterschiedlich: während 1995 im Juli 25 C. acus in 45 S. regalis gefunden wurden; 
waren es 1998 zur selben Jahreszeit nur 12 Fische in 329 S. regalis; 1996, 1997 und 
1999 fischten wir keinen einzigen C. acus. Der Besatz von S. regalis mit C. acus war 
1995 mit 55% viel höher als in allen anderen Jahren, obwohl alle Fänge zur gleichen 
Jahres- und Tageszeit im selben Gebiet stattfanden und auch die angewandten Fang- 
methoden, sowie Boot, Netz und Mannschaft immer die gleichen waren. Die beiden 
einzigen C. acus aus 85 H. tubulosa wurden 1998 in weniger als 10 m Meerestiefe über 
Felsgrund bei Banyuls-sur-mer gefunden. 

Die Stichopus-Exemplare waren aus 70 - 110 m Tiefe geholt worden, weshalb 
alle enthaltenen Fische deutliche Dekompressionsschäden zeigten, die 3 - 4 Stunden 
nachdem sie auf Meereshöhe gebracht worden waren in Form von Gasbläschen auf- 
traten. Im Bereich von Kopf und Magen bildeten sich Gasansammlungen, die manch- 
mal zusätzlich über den ganzen Schwanz verteilt vorkamen. Weil der Auftrieb der 
Fische durch diese im Körper eingelagerten Gase erheblich erhöht wurde, war es ihnen 


338 KERSTIN KLOSS UND WOLFGANG PFEIFFER 


unmöglich im Aquarium zu den Seewalzen am Grund abzutauchen. Zudem wirkten die 
Fische extrem geschwächt. Nur in der Seewalze erholten sie sich; alle C. acus, die es 
nicht schafften sich in eine Seewalze zurückzuziehen, erlagen innerhalb weniger Tage 
ihren Verletzungen. Andere Exemplare mit ähnlich schwerwiegenden Dekompressions- 
schäden erholten sich in der Seewalze binnen maximal 3 Tagen und konnten danach 
auch bei normalem Wasserstand wieder zum After abtauchen um in die Seewalze 
einzudringen. 

Wahlversuche: Da die meisten Individuen von C. acus in S. regalis gefunden 
worden waren, sollte überprüft werden ob S. regalis auch bei einer Wahlmöglichkeit 
bevorzugt wird. Alle Wahlversuche verliefen überraschend gleichförmig. C. acus zeigte 
immer ein ausgesprochenes Interesse an A. tubulosa, was am schnelleren und weiter 
ausholenden Schlängeln seines Schwanzes zu erkennen war. Stieß der Fisch zufällig 
zuerst auf S. regalis, machte er keinerlei Anstalten in sie einzudringen, sondern suchte 
das Aquarium weiter ab. Traf er zuerst auf A. tubulosa, ging er meist sofort in “Lauer- 
stellung”. Diese Stellung ist dadurch gekennzeichnet, daß Kopf und Rumpf regungslos 
bleiben während der Schwanz eine schnelle Schlängelbewegung durchführt. Der 
Körper nimmt dabei vor dem After der Seewalze eine horizontale Lage ein. Aus dieser 
Lauerstellung heraus versucht C. acus in den After der Seewalze zu gelangen sobald 
diese Atemwasser ausstößt. In insgesamt 11 Wahlversuchen mit 3 Fischen drangen alle 
ohne Ausnahme in A. tubulosa ein: Fisch Nr. 30 siebenmal, Nr. B und Nr. 31 je 
zweimal. In den Versuchen mit den Fischen Nr. B und Nr. 31 wurde jeweils einmal 
beobachtet, wie sie in eine schon besetzte A. tubulosa eindrangen, obwohl jedesmal ein 
unbewohnter S. regalis als Alternative zur Verfügung stand. Auch in anderen Ver- 
suchen hielten sich wiederholt 2 - 3 C. acus in einer A. tubulosa oder im selben 
S. regalis auf. Die Wahlversuche zeigten deutlich, daß C. acus H. tubulosa gegenüber 
S. regalis bevorzugt. 

Einstiegsvarianten: C. acus kann auf 2 verschiedene Weisen in die Seewalze 
eindringen - Kopf oder Schwanz voraus. Kopf voraus Eindringen (Abb. 1): C. acus legt 
sich in Lauerstellung vor den After der Seewalze. Sobald sie ihren Atemwasserstrom 
durch den After ausstößt, beschleunigt der Fisch blitzschnell die Schlängelbewegungen 
seines Schwanzes und ist innerhalb von 40-80 ms vollständig in der Seewalze 
verschwunden. Dieser Vorgang ist manchmal so schnell, daß die Einzelbildanalyse 
keinen genauen Aufschlufl ermöglicht: bei S. regalis: oft schneller als 80 ms, manchmal 
aber auch 4, 10, 13, 50 und 65 s; bei H. tubulosa meist zu schnell um mit Aufnahmen 
von 25 Bildern/ s festgehalten werden zu können, oft 40-80 ms, nur ausnahmsweise 
mehrere Sekunden. Schwanz voraus Eindringen (Abb. 2a-c): Wenn C. acus rückwärts 
in die Seewalze einzudringen versucht, nimmt er zuerst dieselbe Lauerstellung ein wie 
beim Vorwärtseindringen. Jetzt ist aber zu beobachten, daß die Schwanzspitze sich 
immer wieder an der rechten oder linken Seite einzurollen und eine Schleife zu bilden 
beginnt. Spürt der Fisch den Wasserstrom der Seewalze, führt er den Schwanz an seiner 
Körperseite und am Kopf vorbei, den Körper gleichsam als Gleitschiene verwendend. 
Der Kopf dient dabei als Einfädelhilfe für den Schwanz, so daß dieser in den geöffneten 
Anus eingeführt werden kann. Die Fische bewegten hierbei den Schwanz in fast allen 
Fällen an ihrer linken Körperseite vorbei. Steckt der Fisch bereits mit der 


BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 339 


ABB. 1. Carapus acus (Länge 115 mm): Vorwärtseindringen in S. regalis; a) Lauerstellung mit 
Maul am After der Seewalze; b) Kopf in Seewalze, kräftiger Schwanzschlag. (Fotos: T. Moritz) 


340 KERSTIN KLOSS UND WOLFGANG PFEIFFER 


Schwanzspitze im After der Seewalze, so biegt er seinen Körper nach einer Seite hoch, 
wodurch ihm anscheinend ein besserer Halt ermöglicht wird. Im Innern der Seewalze 
verkeilt er sich. Je nach Seewalzenart und Fischgröße dauerte es 12 - 472 s bis C. acus 
ganz in der Seewalze verschwunden war: bei S. regalis: 15, 34, 58, 102, 139 und 195 s; 
bei A. tubulosa: 12, 13, 32, 93, 110 und 472 s. Der Rückwärtseintritt währte mehr als 
doppelt so lang wie der Vorwärtseintritt. Unterbrechungen im Ablauf des Eindringens 
fanden an unterschiedlichen Stellen statt. Daß der Fisch dabei durch seine leicht 
abstehenden Brustflossen und Kiemendeckel behindert wird, war nicht zu erkennen. 
Nachdem C. acus ganz in der Seewalze verschwunden war, wurde wiederholt 
beobachtet wie er seinen Kopf durch den After der Seewalze teilweise herausstreckte 
und atmete (Abb. 2d). In manchen Fällen besaßen S. regalis-Exemplare, in welche 
Fische eindrangen, keine weiteren inneren Organe mehr außer Darmresten. Gonade und 
Wasserlunge waren bereits früher ausgeworfen worden. In diesen Fällen ist anzu- 
nehmen, daß der Fisch über dieselbe Öffnung der Enddarmerweiterung in die Leibes- 
höhle vordrang über die er in die Wasserlunge gelangen könnte. Problematisch wird die 
Erklärung allerdings für Fische, die in der Leibeshöhle von Seewalzen gefunden 
wurden, welche noch über ihre Wasserlunge verfügten. Es wird vermutet, daß sich der 
Fisch in der engen Wasserlunge umdreht und dabei das dünnhäutige Lungengewebe 
durchstößt. 

Drei S. regalis wurden beobachtet wie sie auf das Eindringen des Fisches 
umgehend mit heftigem Schlängeln reagierten. Ihre Auf- und Abbewegungen dauerten 
bis zu 8 min und wurden auch durch Umsetzen in ein anderes Aquarium nicht 
unterbrochen; der C. acus im Innern verließ seine Behausung jedoch nicht. A. tubulosa 
versuchte durch Kontraktion ihres Afterschließmuskels und ihrer Hautmuskulatur im 
hinteren Körperabschnitt den Fisch am Eindringen zu hindern. Ihr Körper erschien 
dadurch im posterioren Bereich wesentlich dünner. Wie die lange Einstiegsdauer 
vermuten läßt, wurde dadurch das Vorhaben von C. acus zwar erschwert, doch nicht 
verhindert. In 2 Fällen kotete H. tubulosa, wodurch es ihr gelang den bereits teilweise 
eingedrungenen C. acus wieder herauszudrücken. Beide Male startete der Fisch 
sogleich einen weiteren Versuch, der jeweils schnell zum Erfolg führte. Häufigkeit und 
Verteilung der vor- und rückwärtigen Eintritte von C. acus hängen sowohl von der 
Größe des Fisches als auch von der Seewalzen-Species ab. Bei Fisch Nr. 30 (115 mm) 
halten sich die rück- und vorwärtigen Eintritte in S. regalis mit 9:7 etwa die Waage; bei 
H. tubulosa wurde dagegen der Rückwärtseinstieg mit 20:5 bevorzugt. Zwei weitere 
Fische entschieden sich mit 11:2 bzw. 7:0 ebenfalls für den Riickwartseintritt. Der 
Wechsel zwischen den beiden Eintrittsvarianten erfolgte bei diesen Fischen mittlerer 
Körperlänge unregelmäßig. Größere Fische drangen dagegen immer mit dem Schwanz 
voran ein, egal ob sie S. regalis oder H. tubulosa vor sich hatten. Haut- und Schließ- 
muskulatur sind bei H. tubulosa stärker entwickelt und kräftiger als bei S. regalis, bei 
dem sich der After auch in kontrahiertem Zustand leicht öffnen läßt. Der Fisch kann 
also auch mit seinem dicken Vorderende voraus eindringen. Anders ist die Situation bei 
H. tubulosa. Mit ihrer kräftigen Muskulatur verschließt sie den Anus so fest, daß es 
selbst mit einer dünnen Sonde nahezu unmöglich ist ihn zu öffnen. Hier steckt C. acus 


BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 341 


ABB. 2. Carapus acus (dasselbe Exemplar wie in Abb. 1): (a-c) Rückwärtseindringen in A. 
tubulosa; a) Einrollen der Schwanzspitze; b) Schwanz wird links am Körper vorbeigeführt; c) 
Fisch hat sich umgedreht, Schwanzspitze im After der Seewalze; d) Kopf von oben, in S. regalis; 
e) beim Freipräparieren in S. regalis. (Fotos: T. Moritz) 


342 KERSTIN KLOSS UND WOLFGANG PFEIFFER 


zuerst seinen spitz auslaufenden Schwanz als dünneres Körperende in den Anus. Ist der 
Schwanz erst einmal in der Seewalze, kann den Fisch fast nichts mehr davon abhalten 
ganz einzudringen. Handelt es sich bei dem Fisch um ein größeres Exemplar, 
verwendet er die Variante “Schwanz voraus” auch bei S. regalis, da sein Kopf für einen 
frontalen Vorstoß zu dick ist. 

Mageninhalt: In histologischen Schnittpräparaten von 2 nach Bouin fixierten 
Mägen sind deutlich Komplexaugen von Kleinkrebsen zu erkennen. Dagegen konnten 
histologisch keine Hinweise darauf gefunden werden, daß C. acus auch Gewebe der 
Seewalzen fressen würde. Weder Gonaden noch Muskulatur oder sonstige zelluläre 
Strukturen von S. regalis befanden sich in den Fischmägen. In den Magen von 2 in 
Formalin fixierten Fischen aus S. regalis, wurden 4 Kleinkrebse angetroffen bei denen 
es sich vermutlich um die Garnele Pandalina sp. (Decapoda) handelt (Fischer er al., 
1987). Insgesamt wurden in den Mägen von 4 der 17 untersuchten Männchen und 
Weibchen von C. acus (154 - 195 mm Gesamtlänge) Krebse oder deren Augen 
gefunden. Knapp 1/4 der Fische hatte also kurz vor seinem Fang Krebse gefressen. 


DISKUSSION 


VERBREITUNG: Die meisten Species der Gattung Carapus leben in tropischen 
oder subtropischen, überwiegend seichten Gewässern (Williams, 1984; Markle & 
Olney, 1990). Carapus acus ist auf das Mittelmeer und die angrenzende südliche 
Atlantikküste beschränkt. Arnold (1953, 1956) fand C. acus bei Neapel hauptsächlich 
in H. tubulosa aus 10 - 20 m Meerestiefe. Von 1350 bei Neapel in 10 - 25 m Tiefe 
gesammelten A. tubulosa-Exemplaren enthielten nur 19 (1,4%) einen C. acus (Gustato, 
1976). Der Befall von nur 2 A. tubulosa bei Banyuls-sur-mer hatte den Anschein von 
Zufälligkeit, denn 38 von 40 C. acus fanden wir in S. regalis aus 70 - 110 m Tiefe. Die 
im Juli - August 1998 gefangenen C. acus waren überwiegend Jungfische (Gesamtlänge 
durchschnittlich 129 mm), von denen sich aber keiner mehr in einem Larvenstadium 
befand. Im Gegensatz dazu handelt es sich bei den 1995 zur selben Jahreszeit gefan- 
genen Fischen hauptsächlich um große adulte Exemplare (Gesamtlänge durchschnitt- 
lich 173 mm), wobei der Befall 55% betrug. Im Meer haben wir zwar die meisten (38) 
C. acus in (450) S. regalis und nur 2 in (85) H. tubulosa gefunden, doch als wir die 
Fische im Labor vor die Wahl zwischen S. regalis und H. tubulosa stellten, entschied 
sich C. acus immer für H. tubulosa. Die Bevorzugung von A. tubulosa im Aquarium 
war überraschend, da deren Tiefenverbreitung im Untersuchungsgebiet von Banyuls- 
sur-mer großteils nicht mit der des Fisches übereinstimmt. Bei den zwei in geringer 
Tiefe in A. tubulosa gefangenen C. acus handelte es sich vermutlich um verirrte Indivi- 
duen, obgleich C. acus in anderen Regionen vor allem in A. tubulosa aus geringer Tiefe 
gefischt wurde (Emery, 1880). Der Vergleich der Literaturangaben zur Tiefenver- 
breitung von C. acus zeigt hingegen, daß sich das Vorkommen des Fisches mit dem 
beider Seewalzen-Species in anderen Gebieten überschneidet (Abb. 3). Folgende Hypo- 


these könnte die experimentelle Bevorzugung von H. tubulosa erklären. Geht man 


davon aus, daß sich der Fisch von Kleinkrebsen ernährt, wäre es gleichgültig welche 
Species er als “Wohnhöhle” aussucht. Wichtig wäre dagegen, welches Substrat die See- 


| 


| 


BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 343 


walze bevorzugt, da auch die Beutekrebse untergrundspecifische Vorlieben haben. Die 
als Nahrung nachgewiesene Pandalina sp. lebt überwiegend auf Felsgrund. Auch A. 
tubulosa wurde meistens in Felsnähe gefunden. Als Substratfresser ist diese Seewalze 
zwar auf Sandgrund angewiesen, lebt jedoch vor Banyuls-sur-mer hauptsächlich an 
Orten, an denen die schützende Felsformation der Küste in sandigen Untergrund 
übergeht. Vor Banyuls-sur-mer ist dies in 5 - 20 m Tiefe der Fall. S. regalis kommt 
hingegen vorwiegend auf Sandgrund vor. So ließe sich erklären, warum C. acus H. 
tubulosa gegenüber S. regalis bevorzugt. In der nur bis ca. 25 m tiefen Bucht von Port 
de la Selva mit Sandgrund und Posidonia-Wiesen sind H. tubulosa und besonders 
H. polii zwar häufig, doch fehlen hier C. acus und S. regalis. 

VERHALTEN: Aus einer Seewalze herauspräparierte und im Aquarium frei 
schwimmende C. acus zeigten zwei völlig unterschiedliche Schwimmweisen. Die einen 
schwammen an der Wasseroberfläche mit weit ausholenden schlängelnden Bewegun- 
gen und streckten dabei ihren Kopf immer wieder aus dem Wasser. Die anderen 
tauchten sofort Kopf voran zum Aquarienboden, wobei ihre schräge Schwimmlage 
beibehalten wurde bis sie auf eine Seewalze stießen. Arnold (1953) beschreibt ebenfalls 
diese beiden Schwimmweisen, deutet sie jedoch als Kennzeichen verschiedener Ent- 
wicklungsstadien. Nach Arnold (1953) schwimmen Tenuis-Larven mit weitausholen- 
den Bewegungen während junge und adulte Fische nur leicht mit dem Schwanz 
schlängeln. Unsere Beobachtungen konnten diese Aussage nicht bestätigen. Vielmehr 
hatte es den Anschein, daß die Schwimmweise mit dem Allgemeinbefinden des Fisches 
zusammenhängt. Hatte er Dekompressionsschäden oder war der Sauerstoffgehalt im 
Aquarium zu niedrig, so schnappte er wiederholt an der Oberfläche nach Luft und 
schwamm mit hektischen Bewegungen. Das gleiche Verhalten zeigte er auch in 
Bedrängnis. Dagegen schwamm er bei gutem Allgemeinzustand in Schräglage mit dem 
Maul am Boden. Die beiden Schwimmweisen sind somit nicht abhängig vom 
Entwicklungsstadium sondern Ausdruck des Befindens. Wenn der Fisch vor dem After 
der Seewalze seine Lauerstellung einnimmt, sind die Bewegungen der Brustflossen 
deutlich zu erkennen. Diese Flossen schlagen abwechselnd und dienen sowohl der 
Balance als in geringem Maße auch der Lokomotion. Der Vortrieb erfolgt haupt- 
sächlich über den Schwanz, wie beim Vorwärtseindringen zu sehen ist. Hierbei 
beschleunigt der Fisch in kürzester Zeit bis auf Höchstgeschwindigkeit durch kräftiges, 
weitausholendes Schwanzschlagen. 

Carapus acus findet die Seewalzen vermutlich zufällig; ein besonderes Such- 
schema war nicht zu erkennen. Nach Arnold (1958) und Trott (1981) geschehe die 
Lokalisierung von weiter entfernten Zielen chemisch, während auf kurze Distanz die 
visuelle Orientierung eine Rolle spiele. Van Meter & Ache (1974) vermuten, daß nur 
die Lokalisierung des Afters über die Augen erfolgt. Unsere Fische schwammen im 
Aquarium umher bis sie auf eine Seewalze trafen. Ob und wie weit hierbei chemische, 
taktile oder visuelle Reize eine Rolle spielen, kann nicht entschieden werden. Die 
deutliche Reaktion auf das ausgestoßene Atemwasser ist wahrscheinlich taktil und 
chemisch bedingt. Eine visuelle Wahrnehmung ist nicht anzunehmen, da fast alle 
Versuchsfische aus 70 - 110 m Tiefe stammen und ihre Augen weitgehend rückgebildet 
sind. Die Beobachtung, daß vor allem kleinere Fische vorwärts in die Seewalze 


344 KERSTIN KLOSS UND WOLFGANG PFEIFFER 


eindringen, bestätigt Trott’s (1981) Aussage, daß diese Weise die schnellste Möglich- 
keit des Eindringens darstellt. Die Entscheidung, ob ein Fisch vorwärts oder rückwärts 
eindringt, wird durch die morphologischen Gegebenheiten der Seewalzen-Species und 
die Fischgröße bestimmt, denn auch große Fische können vorwärts und kleine rück- 
warts eindringen. Zweimal wurden am Morgen Fische freischwimmend im Aquarium 
angetroffen, die sich am Vorabend noch in einer Seewalze befunden hatten. Ferner 
entwischten einige Exemplare in der Nacht durch die unzureichend gesicherten über- 
läufe. Dies zeigt, daß C. acus die Seewalze nachts verläßt, vermutlich um auf Beute- 
fang zu gehen. Einmal wurde beobachtet, wie ein C. acus seine H. tubulosa verließ 
nachdem er etwa eine halbe Stunde in ihr verweilt hatte. Der Fisch schwamm einige 
Runden im Aquarıum und verschwand dann wieder in derselben Seewalze. Das 
selbständige Verlassen der Seewalze, ohne von ihr ausgeworfen zu werden, ist also 
möglich. Daß sich C. acus nicht dauernd in der Seewalze aufhält, zeigen auch die im 
Meer freischwimmenden Exemplare (Smith, 1964). 

ERNÄHRUNG: Die anatomischen und histologischen Untersuchungen bewiesen, 
daß C. acus Krebse frißt. Fraglich blieb hierbei, ob diese sein ganzes Nahrungs- 
spektrum ausmachen. Es gibt jedoch keinen Hinweis, daß C. acus seinen Wirt anfrißt. 
In keinem der untersuchten Mägen wurden Reste von S. regalis gefunden. Diese 
Befunde sprechen für die Annahme, daß C. acus kein Nahrungsparasit ist. Während der 
Verhaltensexperimente hatte C. acus nur Seewalzen zur Verfügung von denen er sich 
laut Arnold (1953) parasitisch ernährt. Eine andere potentielle Nahrung wurde in 
unseren Wahlexperimenten nicht angeboten. Auf separate Fütterungsversuche mit 
Artemia sp. oder lebenden Larven von Mysis sp. reagierte C. acus nicht. Wurden die 
Futtertiere ins Aquarium gebracht, so schwamm der Fisch sofort in die dunkelste Ecke 
ohne sich um die Krebse zu kümmern. Pandalina sp., die wir in den Magen von C. acus 
gefunden hatten, war für uns als Futtertier nicht zugänglich. Gab man die Wasserlunge 
von S. regalis oder H. tubulosa ins Aquarium, war wiederholt zu beobachten, daß sich 
der Fisch neben oder unter der Lunge zusammenringelte. Kurzzeitig hatten wir deshalb 
die Coelomocyten der Seewalzen, die auch der Verdauung und Nährstoffspeicherung 
dienen, in Verdacht von C. acus aufgenommen zu werden. In diesem Fall müßten sich 
die Fische trinkend ernähren, indem sie die Coelomflüssigkeit der Seewalze schlucken 
um die in den Coelomocyten gespeicherten Stoffe zu verwerten. Zum einen besitzt 
C. acus keine Strukturen die einen filtrierenden Nahrungserwerb ermöglichen, zum 
anderen müflten gewaltige Mengen von Coelomocyten geschluckt werden um den 
Energiebedarf zu decken. 

Um die Frage eines “Zubrotes” neben Futterkrebsen zu klären, wurde zum 
Vergleich die freilebende Carapinae-Art Echiodon drummondi Thompson, 1837 un- 
tersucht (Trott, 1981). Diese lebt in größeren Tiefen (bis ca. 300 m) vor den Küsten 
Britanniens, des südlichen Norwegens, des westlichen Dänemarks und in der Biskaya 
(Trott & Olney, 1986); sie ernährt sich von kleinen Invertebraten und Fischen. 
Achtzehn Exemplare von E. drummondi, die der Zweit-Autor 1985 vor Arcachon in der 
Biskaya in 200-300 m Meerestiefe gefangen und in Formalin fixiert hatte, wurden auf 
ihren Mageninhalt geprüft. In einem wurde ein vermutlich parasitischer Nematode 
gefunden, 6 andere enthielten Krebse. Wegen der fortgeschrittenen Verdauung waren 3 


BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 345 


100 


120 


400 
AO) FC) 20) Se) eo) mel. 0) 
We bis unterhalb von [ | Hauptverbreitung 
IN bis oberhalb von | È seltenes Vorkommen 


ABB. 3. Vertikale Verbreitung von S. regalis, H. tubulosa und Carapus acus: Vergleich der 
eigenen Befunde (schwarz) mit Literaturangaben (weiß) von: a) Nadal i Fortia (1981, 1994), b) 
Fiedler & Lieder (1994), c) Riedl (1983), d) Mayer (1937), e) Bauchot & Pras (1980), f) Fiedler 
(1991), g) Markle & Olney (1990). Ordinate: Tiefe in m. 


Krebse nicht zu identifizieren, bei den 3 anderen handelte es sich um Isopoda. 
Insgesamt hatte also 1/3 der untersuchten Fische Crustaceen im Magen. Bei C. acus 
betrug der Anteil der Exemplare mit Krebsen im Magen etwa 1/4. Der Vergleich der 
beiden Werte legt die Vermutung nahe, daß auch C. acus kein Nahrungsparasit ist, 
sondern ein nachtaktiver Jäger. Es ist anzunehmen, daß die tagsüber gefischten C. acus 
ihr nächtliches Mahl bereits verdaut hatten. Demgegenüber müßten bei parasitischer 


346 KERSTIN KLOSS UND WOLFGANG PFEIFFER 


Ernährung, die keinem tageszeitlichen Faktor unterliegt, ständig Gonaden oder andere 
Gewebe von Seewalzen in den Fischmägen gefunden werden. Dies war in unserer 
Untersuchung jedoch nicht der Fall. Arnold (1953), der die parasitische Ernährung für 
C. acus postuliert, führt keinen Beweis für seine Spekulation an. Außerdem beobachtete 
Gustato (1976), daß von 17 C. acus, die er aus H. tubulosa entnahm, sich nur 2 in der 
Leibeshöhle befanden. Wie sollten die restlichen 15 Fische in der Wasserlunge von der 
Gonade fressen? 

Um die Hypothese zu untermauern, wonach sich C. acus rein räuberisch ernährt, 
ist ein Vergleich mit den anderen Species der Gattung Carapus hilfreich. Für viele von 
ihnen wird ebenfalls eine räuberische Lebensweise angenommen. Krebse wurden 
gefunden im Magen von C. bermudensis (Smith et al., 1981), C. dubius (Trott, 1970), 
C. homei (Smith, 1964) und C. mourlani (Trott, 1970). Auch C. birpex (Trott & Olney, 
1986) und C. parvipinnis (Fiedler, 1991) werden nicht Parasiten genannt, sondern wie 
die vier vorher aufgelisteten Species als Kommensalen, Einmieter, Inquilinisten, 
Kannibalen oder Symbionten bezeichnet (Strasburg, 1961, Smith & Tyler, 1969; 
Seymour & Mc Cosker, 1970; Dawson, 1971, van Meter & Ache, 1974; Markle & 
Olney, 1980; Fiedler, 1981; Trott, 1981; Trott & Olney, 1986). 

All diese Befunde stützen die Hypothese, wonach auch C. acus als Kommensale 
lebt. Als bessere ökologische Begriffe für die Lebensweise von C. acus können 
Endophoresie (Baer, 1951) und Entökie (Hohorst, 1981; Hentschel & Wagner, 1984) 
herangezogen werden. Am treffendsten erscheint uns die Bezeichnung Inquilinismus 
(Emery, 1880; Bertin & Arambourg, 1958; Trott, 1970; Trott & Olney, 1986), da sich 
der Fisch selbständig ernährt. Völlig fehl am Platz ist hingegen der Begriff Symbiose 
(van Meter & Ache, 1974; Gustato, 1976), da aus unserer Sicht die als Herberge 
dienende Seewalze keinerlei Vorteil von ihrem Mieter hat, der anscheinend allein 
profitiert. Aufgrund Arnold’s (1953) unbewiesener Vermutung ist C. acus bisher zu 
Unrecht als Nahrungsparasit bezeichnet worden. Diese Behauptung wurde von der 
umfangreichen Sekundärliteratur übernommen (Bauchot & Pras, 1980; Riedl, 1983; 
Terofal, 1986; Fiedler, 1991; Fiedler & Lieder, 1994 u.a.), was zur heutigen all- 
gemeinen Verbreitung dieser Ansicht führte. Es gibt jedoch bisher keinen einzigen 
Beweis dafür, daß C. acus von der Seewalze frißt und sich somit parasitisch ernährt. 
Die Seewalze dient ihm vielmehr nur als schützende Zuflucht zwischen seinen nacht- 
lichen Beutezügen. Carapus acus muß dementsprechend als Inquilinist bezeichnet 
werden, wie dies Emery (1880) bereits völlig richtig getan hat - er nennt den Fisch 
“inquilino”! 


SUMMARY 


450 Stichopus regalis (Cuvier), caught with a trawl-net at a depth of 70 - 110 m 
off Banyuls-sur-mer (France), contained 38 pearlfish, Carapus acus (Brünnich). Two 
pearlfish were found in 85 Holothuria tubulosa Gmelin caught at Banyuls-sur-mer in 
less than 10 m of seawater. In contrast, 63 H. tubulosa and 142 H. polii Della Chiaje, 
collected by diving in the bay of Port de la Selva (ca. 10 miles south of Banyuls), did 
not contain any C. acus. Pearlfish collected from 1995 to 1998 were preserved to study 


BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 347 


their stomach contents. Twelve individuals sampled in the summer of 1998 were kept in 
aquaria to analyse their behaviour by means of computer analysis of video records. Due 
to severe damage caused by insufficient decompression only a few individuals survived 
longer than one week after being placed in an aquarium. Since most C. acus were 
collected from S. regalis, the host preference of the fish was tested in an aquarium. 
S. regalis and H. tubulosa were provided as potential hosts. The results were surprising: 
in 11 experiments the 3 fish examined entered only H. tubulosa if they could choose 
between both species. 

The entry of the pearlfish through the anus of the host holothurian may be head 
first or tail first. The pearlfish, in a head-down position, lies in wait in front of a sea 
cucumber. As the sea cucumber discharges its respiratory water the fish rapidly flicks 
its tail and enters the host. In the case of H. tubulosa the pearlfish enters head first 
within 40 - 80 ms; in S. regalis it takes up to 4 - 60 s. Tail first entry of C. acus takes 12 
- 120 s. To enter the host, the pearlfish bends up its tail towards the head and its whole 
body forms a loop. When the waiting pearlfish perceives the water discharged by a 
holothurian, it moves its tail foreward along the side of its body and head. By this, the 
head helps the tip of the tail to target the anus of the host. Three S. regalis responded to 
intruding pearlfish with intense up and down movements; H. tubulosa contracted the 
sphincter of its anus and the spherical muscles of its body wall in the posterior region, 
whereby they became thin. However, neither defence method prevented C. acus from 
entering its host. In C. acus the frequency of head first and tail first entries depended on 
the size of the pearlfish and on the species of the host holothurian. A 145 mm long 
pearlfish entered S. regalis 9 times tail first and 7 times head first. However, the same 
fish entered H. tubulosa 20 times tail first and only 5 times head first. 

The presence of C. acus in S. regalis sampled was extremely variable. In 1995 
as many as 25 C. acus were collected from 45 S. regalis specimens. However, since 
1996 only 13 C. acus were found in 405 S. regalis, although the locality, depth, season, 
day time, boat, trawl-net, sampling method and techniques, and even the crew were the 
same. The C. acus specimens sampled in 1995 were large (0 173 mm) and adult, 
whereas the pearlfish caught in 1998 were much smaller (@ 129 mm) and mostly 
juvenile. 

Pearlfish removed from the host holothurian demonstrated two completely 
different styles of swimming. Either they swam at the water surface with wriggling 
movements, repeatedly raising their head out of the water, or they immediately dived to 
the ground of the tank, maintaining a 30 - 40 degree tilted head-down position until 
they found a host holothurian. The latter style of swimming is only exhibited by healthy 
individuals. Swimming at the surface indicates either damage caused by insufficient 
decompression or a poor physical condition of the pearlfish. 

In the aquarium it was found that pearlfish occasionally leave their host holo- 
thurians, especially at night, when they presumably feed. The stomachs of 4 out of 17 
individuals contained small shrimps, probably Pandalina sp. In comparison, isopods 
and other small crustaceans were found in the stomachs of 6 out of 18 specimens of the 
closely related, but free-living, Echiodon drummondi Thompson. The statement by 
Arnold (1953) that C. acus is a parasite which feeds on the organs (gonads etc.) of its 


348 KERSTIN KLOSS UND WOLFGANG PFEIFFER 


sea cucumber host could not be confirmed. This statement is completely speculative 
and not based on any research results. The stomachs of C. acus examined did not 
contain any holothurian tissue. All other species of the genus Carapus feed on small 
invertebrates and fish, and none feed parasitic. Our results give reason to agree with 
Emery (1880), who stated that C. acus is an inquiline (“inquilino”), to whom the host 
holothurian serves as a shelter only. 


DANKSAGUNG 


Wir danken all denen, die geholfen haben, daß diese Arbeit zustande kam. Unser 
besonderer Dank gebührt der Direktion des Laboratoire Arago in Banyuls-sur-mer 
(Frankreich), die uns ermöglichte, auf diesem Gebiet zu arbeiten. Die Mitarbeiter der 
Station waren immer hilfsbereit. Weder schlechtes Wetter noch stürmische See konnten 
die Mannschaft des Forschungsschiffes Nereis davon abhalten mit uns auszufahren, um 
Seewalzen zu fischen. Besonders zu erwähnen ist unser Kollege Timo Moritz, der 
große Geduld bewies beim photographischen Festhalten des Eindringens von C. acus in 
Seewalzen. 


LITERATUR 


ARNOLD, D. C. 1953. Observations on Carapus acus, (Brunnich) (Jugulares, Carapidae). Pubbli- 
cazioni della Stazione Zoologica di Napoli 24: 152-166. 

ARNOLD, D. C. 1956. A systematic revision of the fishes of the Teleost family Carapidae (Perco- 
morphi, Blennioidea), with descriptions of two new species. The Bulletin of the British 
Museum (Natural History) 4: 247-307. 

ARNOLD, D. C. 1958. Further studies on the behaviour of the fish Carapus acus (Briinnich). 
Pubblicazioni della Stazione Zoologica di Napoli 30: 263-268. 

BAER, J. G. 1951. Ecology of animal parasites (pp. 1-7). The University of Illinois Press, Urbana, 
224 pp. 

BAUCHOT, M. L. & Pras, A. 1980. Guide des poissons marins d’ Europe, pp.196-197. Delachaux 
& Niestlé, Lausanne, 427 pp. 

BERTIN, L. & ARAMBOURG, C. 1958. Famille des Fierasferidae (pp. 2424-2426). In: Grassé, P.-P. 
(ed.). Traité de Zoologie, Tome XIII. Masson et Cie, Editeurs, Libraires de l’Académie 
de Medecine, Paris, 2758 pp. 

Dawson, C. E. 1971. Records of the pearlfish, Carapus bermudensis, in the northern gulf of 
Mexico and of a new host species. Copeia 1971: 730-731. 

EMERY, C. 1880. Le specie del genere Fierasfer nel golfo di Napoli e regioni limitrofe. Fauna 
und Flora des Golfes von Neapel und der angrenzenden Meeres-Abschnitte 2: 1-76. 

FIEDLER, K. 1991. Fische (pp. 188 + 316). Jn: Kaestner, A. Lehrbuch der Speziellen Zoologie Bd. 
II: Wirbeltiere. G. Fischer, Jena, 498 pp. 

FIEDLER, K. & LIEDER, J. 1994. Mikroskopische Anatomie der Wirbellosen. G. Fischer, Stuttgart, 
238 pp. 

FISCHER, W., SCHNEIDER, M. & BAUCHOT, M. L. 1987. Mediterranée et Mer Noire, Vol.1 
Vegetaux et invertébrés (p. 245). CEE et FAO, Rome, 760 pp. 

GUSTATO, G. 1976. Osservazioni sulla biologia e sul comportamento di Carapus acus (Ophi- 
dioidea, Percomorphi). Bolletino della Societa dei Naturalisti in Napoli 85: 505-535. 

HENTSCHEL E. & WAGNER, G. 1984. Zoologisches Wörterbuch. 2. A. (pp. 334-335). G. Fischer, 
Stuttgart, 672 pp. 


BIOLOGIE VON “FIERASFER” (CARAPUS ACUS) 349 


HOHORST, W. 1981. Parasitologie (p. 766). In: Starck, D., Fiedler, K., Harth, P. & Richter, J. 
(eds.), Biologie. Verlag Chemie, Weinheim, 1118 pp. 

KAESTNER, A. 1963. Lehrbuch der Speziellen Zoologie, Teil 1: Wirbellose (pp. 1212-1251). 
G. Fischer, Stuttgart, 1423 pp. 

MARKLE, D. F. & OLNEY, J. E. 1980. A description of the vexillifer larvae of Pyramodon 
ventralis and Snyderidia canina (Pisces, Carapidae) with comments on classification. 
Pacific Science 34: 173-180. 

MARKLE, D. F. & OLNEY, J. E. 1990. Systematics of the pearlfishes (Pisces: Carapidae). Bulletin 
of Marine Science 47: 269-410. 

Mayer, B. 1937. Die Holothurien der Adria, insbesondere der Ktiste von Rovigno. Thalassia 
2: 1-47. 

NADAL I FORTIA, J. 1981. Els nostres peixos. Disputacio de Girona, 255 pp. 

NADAL I Forti, J. 1994. Cataleg dels peixos de la Mediterrània. Anphos, Empuriabrava, 164 pp. 

PFEIFFER, W. 1960. Uber die Schreckreaktion bei Fischen und die Herkunft des Schreckstoffes. 
Zeitschrift fiir vergleichende Physiologie 43: 578-614. 

RIEDL, R. 1983. Fauna und Flora des Mittelmeeres. 3. A. (pp. 588-593 + 712-713). Parey, 
Hamburg, 836 pp. 

SEYMOUR, R. S. & McCosker, J. E. 1974. Oxygen consumption of the commensal fish, Carapus 
homei. Copeia 1974: 971-972. 

SMITH, C. L. 1964. Some pearlfishes from Guam, with notes on their ecology. Pacific Science 18: 
34-40. 

SMITH, C. L. & TYLER, J. C. 1969. Observations on the commensal relationship of the western 
Atlantic pearlfish, Carapus bermudensis, and holothurians. Copeia 1969: 206-208. 

SMITH, C. L., TYLER, J. C. & FEINBERG, M. N. 1981. Population ecology and biology of the 
pearlfish (Carapus bermudensis) in the lagoon at Bimini, Bahamas. Bulletin of Marine 
Science 3: 876-902. 

STRASBURG, D. W. 1961. Larval carapid fishes from Hawaii, with remarks on the ecology of 
adults. Copeia 1961: 478-480. 

TEROFAL, F. 1986. Meeresfische in europäischen Gewässern (pp. 70-71). Mosaik-Verlag, 
Miinchen, 287 pp. 

Trott, L. B. 1970. Contributions to the biology of carapid fishes (Paracanthopterygii: Gadi- 
formes). University of California Publications in Zoology 89: 1-60. 

Trott, L. B. 1981. A general review of the pearlfishes (Pisces, Carapidae). Bulletin of Marine 
Science 31: 623-629. 

Trott, L. B. & OLNEY, J. E. 1986. Carapidae (pp. 1172-1176). In: Whitehead, P. J. P., Bauchot, 
M.-L., Hureau, J.-C., Nielsen, J. & Tortonese, E. (eds.). Fishes of the North-eastern 
Atlantic and the Mediterranean. Vol. 3. UNESCO, Paris, 1473 pp. 

VAN METER, V. B. & ACHE, B. W. 1974. Host location by the pearlfish Carapus bermudensis. 
Marine Biology 26: 379-383. 

WILLIAMS, J. T. 1984. Synopsis and phylogenetic analysis of the pearlfish subfamily Carapinae 
(Pisces: Carapidae). Bulletin of Marine Science 34: 386-397. 


REVUE SUISSE DE ZOOLOGIE, 107 (2): 351-357; juin 2000 


Un nouveau Collembole Neanurinae du Sud du Vietnam, 
Blasconura batäi sp. n., avec une clé des especes du genre 
(Collembola: Neanuridae) 


Anne BEDOS & Louis DEHARVENG 

Laboratoire d’Ecologie Terrestre, UMR 5552 du CNRS, 

U.P.S., 118 route de Narbonne, 31062 Toulouse Cedex 4, France. 
E-mail: deharven @cict.fr 


A new neanurine springtail from southern Vietnam, Blasconura batai 
sp. n., with a key to the species of the genus (Collembola: Neanuridae). 
- The new species is described from a karstic area in southern Vietnam. A 
checklist and a key of all species of the genus Blasconura are given. 


Key-words: Collembola - Neanuridae - Vietnam. 


INTRODUCTION 


Les collines de Hon Chong, isolées en bordure du Golfe de Thailande dans le 
Sud du Vietnam, constituent un remarquable site karstique dont une grande partie a été 
récemment détruite par l’extension de carrieres pour l’exploitation du calcaire. Nous 
avons pu y réaliser quelques récoltes qui nous ont permis de décrire deux especes de 
Collemboles, Lepidonella lecongkieti Deharveng & Bedos, 1995 et Rambutsinella 
honchongensis Deharveng & Bedos, 1996. Nous décrivons dans le présent travail une 
troisieme espece de cette région, Blasconura batai sp. n., et proposons une clé des 
especes du genre Blasconura. 


Blasconura Cassagnau, 1983 


Le genre Blasconura a été créé pour l’espece Achorutes hirtellus Boerner, 1906 
de Java. La description originale de cette espèce, très sommaire, peut s’appliquer à de 
nombreux Neanurinae tropicaux de la tribu des Paleonurini. Il n’est donc pas étonnant, 
étant donné la confusion systématique qui régnait au sein des Neanurinae avant les 
mises au point de Cassagnau (1983, 1986, 1989), que l’espèce hirtella ait été signalée 
dans de nombreuses régions d’Asie du Sud-Est jusqu’en Australasie. Cette vaste distri- 
bution concerne probablement des espèces différentes, comme en témoignent les illus- 
trations données dans la littérature (Handschin, 1926; Womersley, 1933, 1936; Denis, 


Manuscrit accepté le 18.11.1999 


32 ANNE BEDOS & LOUIS DEHARVENG 


1948; Yosu, 1959, 1966; Gapud, 1968). Aucune de ces formes ne peut être attribuée 
sans ambiguïté au véritable B. hirtella tel que l’a redécrit Yosii (1976) d’après un 
matériel topotypique de Bogor (Java), ce que confirment nos propres observations sur 
de nouveaux exemplaires provenant de la localité-type. Autrement dit, toutes les 
citations de B. hirtella antérieures au travail de Yosu (1976) devront être révisées. 

En 1988, Cassagnau décrit 6 Blasconura nouveaux d’Inde et de Sri Lanka; il 
suggere alors de placer Achorutes separatus Denis, 1934 du Vietnam dans le genre 
Blasconura. En suivant cette conception, ce sont 9 especes qui peuvent étre rapportées 
ace genre: 

- Blasconura hirtella (Boerner, 1906) sensu Yosii (1976) 

= Achorutes hirtellus Boerner, 1906 

= Vitronura hirtella; Y osii (1976), Yoshii (1981), Yoshii & Suhardjono (1989) 

= Blasconura hirtella, Cassagnau (1983, 1988) 

= Vitronura (Blasconura) hirtella; Yoshi & Suhardjono (1992) 
espèce décrite de Java et citée d’autres îles indonésiennes (Halmahera et Kei, Yoshii & 
Suhardjono, 1992), de Thailande et de Singapour (Yosu, 1976), et de Malaisie (Sabah, 
Yoshii, 1981). Les citations du Vietnam et des Philippines reprises par Yoshii & 
Suhardjono (1989) d’apres d’anciennes descriptions de Denis (1948) et Gapud (1968) 
sont à vérifier (cf. ante). La citation de Taiwan (Yoshii, 1981) demande également 
confirmation. 

- B. separata (Denis, 1934) comb. n. (Vietnam) 

= Achorutes separatus Denis, 1934 
- B. anamalensis Cassagnau, 1988 (Inde) 

- B. ceylonica Cassagnau, 1988 (Sri Lanka) 
- B. palniensis Cassagnau, 1988 (Inde) 

- B. prabhooi Cassagnau, 1988 (Inde) 

- B. sholica Cassagnau, 1988 (Inde) 

- B. toda Cassagnau, 1988 (Inde) 

- B. batai sp. n. (Vietnam). 


Clé des especes du genre Blasconura 


| Tubercules antennaires soudés entre eux et soudés au frontal en une seule 

MASSE e Up D) 
- Tubercules antennaires soudés entre eux mais séparés du frontal; griffe 

Sansidenty st). dele seh gun. BADER ARE LM er ERBOSO O SE 7 
2 Soie O absente sur le tubercule antenno-frontal; griffe avec une dent........ 3 
- Sole ©) présente sur leitubereule'antenno-frontal «teens 5 
3 Ant. I avec 7 soies; macrochètes écailleux; formule des Di: 122/22222 

PE a EN CAL HE CL TESTE batai sp. n. 

Ant.l'avec'9soles.. 02:45 48e 0 ME NN EEE 4 
+ Macrochètes plumeux formule des Diz 133/22223) eee hirtella 


- Macrochètes longs et lisses; formule des D:244/53553 eee toda 


NOUVEAU COLLEMBOLE NEANURINAE DU VIETNAM 353 


5 Griffe sans dent; ant. I avec 8 soies; macrochètes raides, légèrement 

HUCUCUK: TOFMULE GES D1:4193/22222 Dir. n ceylonica 
- Griffe avec une forte dent; ant. I avec 7 soies; formule des Di: 134/44433 . . . . 6 
6 Macrochetes longs, fortement barbelés dans le tiers distal ............: sholica 
- Macrochetes’eountsstaiblementecalleur un... me ee prabhooi 
7 Soie O absente sur la tête; ant. I avec 7 soies; macrochetes faiblement 

nuoueux-ormule des D0122/2292 RR E ee anamalensis 
- SOI OMPLESC MCS ET N er A celine OSARE 8 
8 Soie O sur les tubercules antennaires soudés; un tubercule supplémen- 

taire entre Di et De sur th. II a abd. III; macrochetes fortement barbelés et 

renflés à l’apex; formule des Di: 233/33333; ant. I avec 9 soies ..... palniensis 
- Soie O sur le tubercule frontal; pas de tubercule supplémentaire sur th. II 

a abd. HI; macrochetes ciliés; formule des Di: 333/22222; nombre de 

solesssurantl-ineonnu es ae os eee ns Pepe Re ae vee separata 


Blasconura batai sp. n. 


MATERIEL TYPE. Holotype femelle et 4 paratypes (2 femelles, 1 mâle subadulte et 1 
juvénile) en préparation microscopique, | paratype juvénile en alcool. Vietnam: province de Kien 
Giang, Hon Chong, colline calcaire de Ba Tai, Berlese de litiere dans un maquis, 19/12/1994 
(relevé VIET-034). Holotype et 3 paratypes déposés au Laboratoire d’Ecologie Terrestre 
(Université Paul Sabatier, Toulouse, France), 2 paratypes déposés au Muséum d’histoire naturelle 
de Geneve (Suisse). 

Les abréviations utilisées dans la description sont celles définies par Deharveng 
(1983) et Deharveng & Weiner (1984). Le matériel étudié a été récolté par les auteurs. 

DESCRIPTION: Longueur: 0,84 a 1,13 mm. Couleur: rouge sur le vivant, blanche 
en alcool, yeux non pigmentés. Habitus assez trapu et convexe (fig. 1). Tubercules 
dorsaux bien délimités mais peu convexes; grain tertiaire marqué par des groupes de 
grains secondaires non soulignés par de véritables réticulations. Abd. VI bilobé, visible 
dorsalement. Soies ordinaires dorsales de 2 types: quelques soies fines, lisses et 
pointues (mésochètes ou microchetes); macrochetes plus ou moins longs, courbes, 
élargis dans leur partie distale en forme de massue, avec une ciliature courte, serrée et 
imbriquée, en forme de pousse d’asperge (fig. 2). Soies S fines, atteignant 1/3 (sur le 
thorax) a 1/2 (a l’arrière de l’abdomen) de la longueur du macrochète voisin. 

Tête (fig. 1, tab Ia et b). Soies S d’ant. IV de taille moyenne, subégales, relati- 
vement fines; vésicule apicale nettement trilobée. Ant. I avec 7 soies. Cône buccal fort; 
labre ogival assez large, avec 2/2,2 soies (les soies distales longues, dépassant l’apex du 
labre). Maxille styliforme, mandibule fine à 3 dents. 2+2 cornéules non pigmentées, la 
cornéule antérieure est extérieure et en avant du tubercule oculaire. Tubercules An et Fr 
fusionnés, séparés de Oc et de CL. Tubercules Di et De faiblement différenciés, non 
fusionnés. Tubercules DL, L et So soudés. Soies O et E absentes. Microchètes Di2 et 
De2 sur la bordure interne du tubercule De. 

Thorax et abdomen. Tuberculisation et chétotaxie illustrées sur la figure | et le 
tableau Ic. Sur abd. IV, le macrochète Di2 est deux fois plus court que le macrochéte 
Dil. Dent interne présente sur la griffe. Soie M absente sur le tibiotarse. 


354 ANNE BEDOS & LOUIS DEHARVENG 


Fics 1-2. Blasconura batai sp. n. 1: habitus et chétotaxie dorsale (longueur du corps: 0,94 mm); 
une soie surnuméraire est présente sur le tubercule De de th. II (côté droit); 2: macrochete Del 
d’abd. I (longueur de la soie: 73 um). 


NOUVEAU COLLEMBOLE NEANURINAE DU VIETNAM 355 


REMARQUE: L’holotype présente une anomalie sur th. II (fig. 1). Les 4 paratypes 
examinés sont également asymétriques pour la chetotaxie d’au moins un de leurs 
tergites. Une forte proportion d’exemplaires non symetriques n’est pas rare chez 
certains Neanurinae de la tribu des Lobellini, mais reste exceptionnelle chez les 
Paleonurini. 

DERIVATIO NOMINIS: Le nom de l’espece est celui de la localité-type, la colline 
de Ba Tai, une des rares collines du karst de Hon Chong qui devrait être épargnée par 
les carrieres de calcaire. 


TABLEAU Ia - Chétotaxie céphalique dorsale de Blasconura batai sp. n. 


Tubercule Nombre Type de Soies 


de soies soles 
CL + 4 ML F 
mé G 
Af + 8 ML A,B 
mi ou mé C,D 
Oc + 3 ML Ocm, Ocp 
mi Oca 
Di + 1 ML Dil 
De + 3 ML Del 
mi Di2, De2 
DL+L+So + 15 ML 4 soies 
Mc 1 soie 
mé 7 soies 


mi 3 soles 


TABLEAU Ib - Chétotaxie céphalique ventrale de B. batai sp. n. 


Vi 6 

Ve 5-6 
Labrum 2129) 
Labium 10,0x? 
Ant. I-II 7,11 
Ant. III 1745S 


Ant. IV or+8S+i+?mou 


Di De DL L Scx2 Cx Tr Fé Ti 
Th. I 1 2 1 - 0 3 6 ? 18 
Th. I 2. 3+S 3+S+ms 3 2 7 Ÿ ? 18 
Th. II À) 4+S 3+S 3 À) 8 ? 11 17 
Abd. I 2 3+S 2 3 TV=4+4 
Abd. II 2 3+S 2 3 Ve=3-4 (Vel absent) 
Abd. III 2 3+S 2 4 Fu=4,0mi Ve=3-4 
Abd. IV 2 2+S 3 6 Ve=7 VL=5 
Abd. V 2 ——3+S$— (3)-4 Ag=3-(4)+3-(4) VL avec L 


Abd. VE —j7 — Ve=14 An=2-3 


356 ANNE BEDOS & LOUIS DEHARVENG 


DISCUSSION 
Deux especes de Blasconura étaient déja connues du Vietnam: 


- Blasconura sp., signalée sous le nom de Achorutes hirtellus Boerner, 1906 du “Centre 
Annam”, a Bana près de Tourane (Denis, 1934, 1948); il s’agit probablement d’une 
espece inedite qui differe du vrai B. hirtella de Java notamment par la présence de 3+S 
soies De (au lieu de 2+S) sur abd. IV. 


- B. separata (Denis, 1934), signalée sous le nom de Achorutes separatus Denis, 1934 
du “Tonkin”, province de Phu-Ho, pres de Yenbai (Denis, 1934, 1948). 


Blasconura batai sp. n. est proche de la première de ces espèces mais possède 
une chétotaxie dorsale nettement plus réduite. La nouvelle espece partage avec B. 
hirtella et B. toda les caractères suivants: la soudure des tubercules An et Fr, l’absence 
de soie O sur la tête et la présence d’une dent à la griffe: B. batai sp. n. se distingue de 
ces deux especes par plusieurs caracteres chétotaxiques, en particulier la présence de 7 
soies sur ant. I au lieu de 9. 


REMERCIEMENTS 


La campagne de récoltes 1994 a Hon Chong a été possible grâce a l’aide de nos 
collègues du département d’Ecologie de l'Université de Ho Chi Minh-Ville et du 
Service Science, Technologie et Environnement de la province de Kien Giang, que 
nous tenons à remercier. 


RÉFÉRENCES 


CASSAGNAU, P. 1983. Un nouveau modèle phylogénétique chez les Collemboles Neanurinae. 
Nouvelle Revue d’Entomologie 13 (1): 3-27. 


CASSAGNAU, P. 1986. Sur l’évolution des Neanurinae paucituberculés à pièces buccales réduites 
(Collemboles) (pp. 313-317). Jn: Dallai, R. (ed.). 2nd International Seminar on Aptery- 
gota, Siena. 

CASSAGNAU, P. 1988. Les Collemboles Neanurinae des massifs du sud de l’Inde et de Ceylan. 
Travaux du Laboratorie d’Ecobiologie des Arthropodes Edaphiques, Toulouse 5 (4): 
2151. 

CASSAGNAU, P. 1989. Les Collemboles Neanurinae: éléments pour une synthese phylogénétique 
et biogéographique (pp. 171-182). Jn: Dallai, R. (ed.). 3rd International Seminar on 
Apterygota, Siena. 

DEHARVENG, L. 1983. Morphologie évolutive des Collemboles Neanurinae, en particulier de la 
lignée Néanurienne. Travaux du Laboratoire d’Ecobiologie des Arthropodes Edaphiques, 
Toulouse 4 (2): 1-63. 

DEHARVENG, L. & WEINER, W. 1984. Collemboles de Corée du Nord. III - Morulinae et Neanu- 
rinae. Travaux du Laboratoire d’Ecobiologie des Arthropodes Edaphiques, Toulouse 4 
(4): 1-61. 

DENIS, J. R. 1934. Collemboles d’Indochine récoltés par M. C. N. Dawidoff (lere note 
préliminaire) [Achorutini]. Bulletin de la Société Entomologique de France 8: 117-122. 

DENIS, J. R. 1948. Collemboles d’Indochine. Récoltes de M. C. Dawydoff. Notes d’Entomologie 
chinoire 12 (17): 183-311. 

Gapub, V. P. 1968. Studies on Philippine Collembola I. Suborder Neoathropleona. The Phi- 
lippine Entomologist 1 (2): 73-96. 


NOUVEAU COLLEMBOLE NEANURINAE DU VIETNAM 357 


HANDSCHIN, E. 1926. Ost-indische Collembolen, IH, Beitrag zur Collembolenfauna von Java und 
Sumatra. Treubia 8: 446-461. 

WOMERSLEY, H. 1933. A preliminary account of the Collembola Arthropleona of Australia. I. 
Poduroidea. Transactions of the Royal Society of South Australia 57: 48-71. 

WOMERSLEY, H. 1936. Further records and descriptions of Australian Collembola. Records of the 
South Australian Museum 5: 475-485. 

YosHu, R. 1981. Neanurid Collembola of Sabah. Entomological Report from the Sabah forest 
research centre 4: 52-68. 

YOSHII, R. & SUHARDJONO, Y. R. 1989. Notes on the Collembolan Fauna of Indonesia and its 
vicinities. I. Miscellaneous notes, with special references to Seirini and Lepidocyrtini. 
Acta Zoologica Asiae Orientalis 1: 23-90. 

YosH, R. & SUHARDJONO, Y. R. 1992. Notes on the Collembolan Fauna of Indonesia and its 
vicinities. II: Collembola of Irian Jaya and Maluku Islands. Acta Zoologica Asiae 
Orientalis 2: 1-52. 

Yost, R. 1959. Studies on the Collembolan fauna of Malay and Singapore. Contributions from 
the Biological Laboratory Kyoto University 10: 1-65. 

Yost, R. 1966. Neanurid Collembola of Australia preserved in the South Australian Museum. 
Records of the South Australian Museum 15 (2): 261-274. 

Yosn, R. 1976. On some Neanurid Collembola of Southeast Asia. Nature and Life in Southeast 
Asia 7: 257-299. 


REVUE SUISSE DE ZOOLOGIE 107 (2): 359-388; juin 2000 


Die Leucospidae (Hymenoptera: Chalcidoidea) der Schweiz, 
mit einem Bestimmungsschlüssel und Daten zu den 
europäischen Arten 


Hannes BAUR! & Felix AMIET? 

! Abteilung Wirbellose Tiere, Naturhistorisches Museum, 
Bernastrasse 15, CH-3005 Bern, Schweiz. 
E-mail: hannes.baur@nmbe.unibe.ch 

2 Forststrasse 7, CH-4500 Solothurn, Schweiz. 


The Leucospidae (Hymenoptera: Chalcidoidea) of Switzerland, with a 
key and data on the European species. - In this paper the taxonomy and 
distribution of the Swiss species of Leucospis Fabricius are presented. Four 
of the seven European species are recorded for Switzerland: L. biguetina 
Jurine, L. dorsigera Fabricius, L. gigas Fabricius, and L. intermedia Illiger. 
Except for L. dorsigera the main distribution lies in the Southern parts of 
the country. The altitudinal gradient ranges from the colline to the 
subalpine belt. Adults appear from May to September. L. biguetina and L. 
gigas are considered vulnerable with regard to the Swiss fauna. Almost 
exclusively females of L. biguetina and L. gigas are known from the study 
area, hence these species are likely to reproduce by thelytoky. A key, 
diagnoses, morphometric and distributional data are also given for the other 
two European species, L. bifasciata Klug and L. brevicauda Fabricius. The 
hosts of all species are critically reviewed and new hosts are recorded for 
L. dorsigera and L. gigas. 


Key-words: Leucospis - taxonomy - key - distribution - new host records - 
Switzerland - Europe. 


EINLEITUNG 


Die Leucospidae stellen mit einer Grösse von ca. 4-16 mm die auffallendsten 
Vertreter der zumeist winzigen Erzwespen (Chalcidoidea) dar. Die Familie ist 
weltweit mit ca. 140 Arten in 4 Gattungen verbreitet (Boutek, 1974; Noyes, 1998), 
wobei in Europa nur gerade 7 Arten der Gattung Leucospis Fabricius vorkommen. 
Durch die verdickten Hinterbeine, den über den Gaster nach vorne gebogenen 
Legebohrer und die gelb-schwarze Zeichnung sind sie unverkennbar. Alle euro- 
päischen Arten führen eine parasitische Lebensweise, d. h. die Larven entwickeln sich 
als Ectoparasitoide an Larvenstadien von Wildbienen, z. B. Anthidium, Megachile 
oder Osmia (Apidae: Megachilinae) (Clausen, 1940; Bouéek, 1974). 

Erste Hinweise auf die Leucospiden der Schweiz finden sich bereits in Sulzer’s 
Geschichte der Insekten (1776), wo unter der treffenden Bezeichnung 'Schenkel- 
wespe’ nicht wie angegeben L. dorsigera sondern L. gigas aufgrund von Exemplaren 


Manuskript angenommen 25.01.2000 


360 HANNES BAUR & FELIX AMIET 


TABELLE | 
Morphologische Terminologie, sortiert nach Abkürzung bzw. Begriff (vgl. Abb. la-Id, gegen- 


überliegende Seite). 


Ag Auge Auge Ag 
Cly Clypeus Clypeus Cly 
Col Procoxa Dorsellum Do 
Co2 Mesocoxa Gastraltergum Gt 
Co3 Metacoxa Mesepisternum Mep 
Do Dorsellum Mesocoxa Co2 
Fel Profemur Mesofemur Fe2 
Fe2 Mesofemur Mesoscutum Msc 
Fe3 Metafemur Mesotarsus Ta2 
Gt Gastraltergum Mesotibia TH 
Mep Mesepisternum Metacoxa Co3 
Mpl Metapleuron Metafemur Fe3 
Msc Mesoscutum Metapleuron Mp1 
Pe Pedicel Metatarsus Ta3 
Ppd Propodeum Metatibia Ti3 
Prn Pronotum Pedicel Pe 
Sca Scapus Procoxa Col 
Scr Scroben Profemur Fel 
Sct Scutellum Pronotum Prn 
Tal Protarsus Propodeum Ppd 
Ta2 Mesotarsus Protarsus Tal 
Ta3 Metatarsus Protibia Til 
Til Protibia Scapus Sca 
Ti2 Mesotibia Scroben Ser 
Ti3 Metatibia Scutellum Sct 


aus Genf beschrieben und illustriert wird. Fuessly (1783) meldet in seinem Beitrag 
zur Lebensweise von ‘L. dorsigera’ ebenfalls Funde von L. gigas aus Genf und der 
Gegend von Zürich. Etwas später beschreibt Jurine (1807) in seinen «Nouvelle 
méthode de classer les hyménopteres et les dipteres» neu L. biguetina Jurine aus der 
Schweiz (vgl. Boutek, 1974), daneben erwähnt er auch die Arten L. dorsigera und L. 
gigas. Hagenbach (1822) stellt aus der Umgebung von Basel L. fuesslini Hagenbach 
auf, ein Synonym von L. dorsigera (vgl. Bouéek, 1974). Labram & Imhoff (1836) 
beschreiben und illustrieren letztere und fügen an, dass in der Schweiz noch 2-3 
weitere Arten vorkämen. Von Chevrier (1870) erhalten wir unter L. lepida Chevrier 
eine weitere Fundortsangabe von L. dorsigera (vgl. Bouëek, 1974) aus dem Genfer- 
seegebiet. Schliesslich zählt Schletterer (1890) in seiner bedeutenden, weltweiten 
Revision der Leucospiden eine Reihe neuer Funde für die erwähnten Arten ın der 
Schweiz auf und führt zusätzlich L. intermedia für das Wallis an. Spätere Arbeiten (z. 
B. Boutek, 1959, 1974; Madl, 1990; Schmiedeknecht, 1930) enthalten kaum neue 
Angaben und zitieren zumeist in allgemeiner Form aus älteren Schriften. 

Während wir recht gut über die Fauna vieler europäischer Ländern unterrichtet 
sind (Boucek, 1959, 1964, 1970, 1977; Berland, 1934b; Madl, 1989, 1990; Pagliano, 
1998; Schletterer, 1890; Schmid-Egger, 1995), fehlte es bislang an einer 
übersichtlichen Darstellung der Leucospiden der Schweiz. Zudem förderte allein die 
grobe Durchsicht der Sammlungen von wenigen Museen ein reiches, unpubliziertes 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 36] 


Kopfbreite 


Metafemurlänge 


Flagellum 


1c 


Gaster 


Ovipositorscheiden 


ABB. la-d, L. gigas © : Kopf frontal (a), Metafemur lateral (b), Antenne lateral (c), Körper ohne 
Kopf lateral (d) (vgl. Tab. 1, gegenüberliegende Seite). Massstab | mm. 


Material aus dem Gebiet zutage. Mit der vorliegenden Arbeit wird erstmals die 
Verbreitung der Arten in der Schweiz umfassend dargestellt. Hierzu wurde das 
Material von sämtlichen grösseren Sammlungen der Schweiz und mehreren Museen 
im Ausland systematisch aufgearbeitet. Die Daten wurden im Hinblick auf eine 
Gefährdung der Arten in der Schweiz ausgewertet. Aufgrund von Funddaten der 
Wirte wurde versucht, den Grad der Übereinstimmung in der Verbreitung von Wirt 
und Parasitoid in der Schweiz aufzuzeigen. Ferner wurde für alle europäischen Arten 
die Literatur nach Wirtsangaben durchgesehen, welche kritisch analysiert und durch 


362 HANNES BAUR & FELIX AMIET 


neue Daten ergänzt wurden. Schliesslich werden ein Bestimmungsschlüssel, 
Diagnosen und morphometrische Daten zu allen europäischen Arten vorgelegt. 


MATERIAL UND METHODEN 


Die vorliegende Studie wertet Daten von insgesamt 919 Exemplaren aus der 
Schweiz (572) und Teilen von Europa, dem Nahen Osten und Nordafrika (347) aus. 
Für die Verbreitungskarten und die Diagramme zur Höhenverbreitung sowie Phäno- 
logie wurde ausschliesslich Material aus der Schweiz verwendet. Den Diagnosen und 
Messungen liegt das bei den einzelnen Arten aufgelistete Material zu Grunde. Die 
betreffenden Exemplare wurden mit einer Etikette mit dem Vermerk «Baur» und 
einer Seriennummer versehen. Messungen wurden an getrocknetem Material unter 
einem Stereomikroskop (Leica MZ12) bei 6 (Körperlänge) bzw. 20facher (alle 
übrigen Masse) Vergrösserung mit Hilfe eines geeichten Okular-Mikrometers (12 mm 
unterteilt in 120 Einheiten) durchgeführt, anschliessend wurden die Werte in Milli- 
meter umgerechnet. Die Körperlänge entspricht der Länge vom Vertex bis zur Spitze 
des Gasters. Diese Messungen können systematische Effekte (Fehler) aufweisen, 
welche von den unterschiedlichen Konservierungsmethoden der einzelnen Sammler 
(z. B. die Streckung bzw. Krümmung von Individuen in einer bestimmten Weise) 
herrühren. Die Kopfbreite, die Metafemur- und die Ovipositorscheidelänge sind frei 
von systematischen Effekten. Die betreffenden Messstrecken sind aus den Abbil- 
dungen la, Ib und Id ersichtlich. Der Beginn der Ovipositorscheide (vgl. Abb. 1d) ist 
gekennzeichnet durch eine feine Kerbe auf der dorsalen Seite des Ovipositors. Meta- 
femur und Ovipositorscheide der linken Seite wurden vermessen. In der morpho- 
logischen Terminologie folgen wir Gibson (1997), die verwendeten Begriffe werden 
in den Abb. la-1d erläutert (vgl. Tab. 1). Quantitative Angaben beziehen sich auf die 
Gesamtheit des betreffenden Materials und sind entsprechend dem System von Goulet 
& Mason (1993: 64) wie folgt definiert: sehr selten <1%, selten 1-10%, gelegentlich 
>10-25%, manchmal >25-50%, oft >50-75%, häufig >75-90%, meist >90-99%, nor- 
malerweise >99% des vorliegenden Materials. 

Eine Beurteilung der Gefährdung der einzelnen Arten in der Schweiz muss bei 
Parasitoiden, besonders bei streng oligophagen Arten wie Leucospis, naturgemäss die 
Gefährdung der Wirte berücksichtigen. Die Angaben richten sich deshalb weitgehend 
nach der Roten Liste der gefährdeten Bienen der Schweiz (Amiet, 1994). Zusätzlich 
wurde der Anteil rezenter Daten (Funde nach 1950) gewertet. Der Beginn für rezente 
Daten wurde auf einen relativ frühen Zeitpunkt festgelegt, da in den letzten drei 
Jahrzehnten die Sammeltätigkeit betreffend Leucospiden stark abnahm und sich mehr 
oder weniger auf die Anstrengungen von FA beschränkte! Die normalerweise geringe 
Populationsdichte sowie die spärliche Besammlung erschweren generell eine Ein- 
schätzung des eigentlichen Bestandes. Wenn jedoch eine Art früher bedeutend 
häufiger festgestellt wurde als in jüngster Zeit (nach 1950), so kann ein Bestan- 
desrückgang angenommen werden (vgl. L. gigas). 

Die Daten für das Verbreitungsbild der Wirte stützen sich auf Erhebungen von 
FA in folgenden Sammlungen: ETHZ, FA, NMBA, und NMBE. Die Nomenklatur der 
mitteleuropäischen Bienen folgt der Liste von Westrich & Dathe (1997). Alle anderen 
Namen werden nach Schwarz er al. (1996) zitiert. Die Namen von Blütenpflanzen 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 363 


richten sich nach der «Flora Helvetica» von Lauber & Wagner (1996). 

Für die Auswertung der Messreihen bzw. die Erstellung der Verbreitungs- 
karten, Grafiken und Abbildungen fand die folgende Software Verwendung: DMAP, 
Version 6.5, 1998 (Verbreitungskarten), Adobe Photoshop, Version 5.0.2, 1998 
(Beschriftung und Nachbearbeitung der Abbildungen und einiger Diagramme), SPSS, 
Version 9.0, 1998 (Auswertung, Diagramme). 

Schliesslich sei für alle weiterführenden Angaben zur Synonymie, Morpho- 
logie und Literatur der behandelten Arten auf die Monographie der Leucospiden von 
Bouéek (1974) sowie den CD-ROM Katalog von Noyes (1998) verwiesen. 


Abkürzungen der Kantone der Schweiz 


BE Bern so Solothurn 

GE Genf TI Tessın 

GR Graubünden VD Waadt 

NE Neuenburg VS Wallis 

SH Schaffhausen ZH Zürich 

Abkürzungen der Sammlungen 

AS Alı Salvioni, Privatsammlung, Zürich, Schweiz 

ASE Albert Sermet, Privatsammlung, Yverdon, Schweiz 

BZOL  Biologie-Zentrum des Oberösterreichischen Landesmuseums, Linz, 
Österreich 

(ES) Christian Schmid-Egger, Privatsammlung, Maulburg, Deutschland 

DEI Deutsches Entomologisches Institut, Eberswalde, Deutschland 

ES Erwin Steinmann, Privatsammlung, Chur, Schweiz 


ETHZ Eidgenössische Technische Hochschule, Entomologische Sammlung, 
Zürich, Schweiz 


FA Felix Amiet, Privatsammlung, Solothurn, Schweiz 

FB Frank Burger, Privatsammlung, Orlamünde, Deutschland 

FNSM  Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt a. M., 
Deutschland 

GC Gilles Carron, Privatsammlung, Neuchätel, Schweiz 

GR Gerd Reder, Privatsammlung, Flörsheim-Dalsheim, Deutschland 

HB Heinrich Bürgis, Privatsammlung, Worms, Deutschland 

HT Hansueli Tinner-Guler, Privatsammlung, Landquart, Schweiz 

TW Ingmar Wall, Privatsammlung, Mihlingen, Deutschland 

MH Martin Hauser, Privatsammlung, Urbana, Illinois, USA [Material 
7 Thy (ESS) 


MHNG Muséum d’histoire naturelle, Genève, Schweiz 

MNHU Museum für Naturkunde, Humboldt-Universität, Berlin, Deutschland 
MSNV Museo Civico di Storia Naturale, Venezia, Italien 

MZL Musée Zoologique, Lausanne, Schweiz 

NMBA  Naturhistorisches Museum, Basel, Schweiz 

NMBE  Naturhistorisches Museum, Bern, Schweiz 

NMSO Naturmuseum, Solothurn, Schweiz 

NMW  Naturhistorisches Museum, Wien, Österreich 


364 HANNES BAUR & FELIX AMIET 


SMNS _ Staatliches Museum für Naturkunde, Stuttgart, Deutschland 


WA Werner Arens, Privatsammlung, Bayreuth, Deutschland 
WL Walter Linsenmaier, Privatsammlung, Ebikon, Schweiz 
ZSM Zoologische Staatssammlung, München, Deutschland 


Bestimmungsschlüssel für die europäischen Arten von.Leucospis (Massstab der 
Abbildungen | mm).! 


1)— Metatibia dorsal rotbraun bis schwarz, höchstens in der Mitte mit 
feinem gelblichem Längsstreifen; basaler Zahn des Metafemurs unge- 
fähr so lang wie folgende Zähne (Abb. 2a). d: Gaster im basalen 
Drittel stark verschmälert (Abb. 2b); 2: Ovipositorscheide sehr kurz, 

nur 0.66-0.74 mal so lang wie der Metafemur............... L. brevicauda 
—  Metatibia dorsal gelb; basaler Zahn des Metafemurs kürzer oder länger 
als folgende Zähne (Abb. 2a', 2a"). d : Gaster im basalen Drittel breiter 
(Abb. 2b'); 2: Ovipositorscheide deutlich länger, 0.90-1.93 mal so lang 


wieder Metatemüur. u... ee ee LO RR 2 
2)—  Basaler Zahn des Metafemurs kürzer als folgende Zähne (Abb. 2a'). ....... 3 
—  Basaler Zahn des Metafemurs länger als folgende Zähne (Abb. 2a") ....... 4 


\ 


2b N 2b' 


' Caleca et al. (1995) geben L. miniata Klug für die kleine Insel Lampedusa (Italien) 
an. Die vor allem in Nordafrika und dem Nahen Osten verbreitete Art (BouCek, 1974) wurde 
im Schlüssel nicht berücksichtigt. L. miniata ist im Habitus sehr ähnlich der orangeroten Form 
von L. gigas, sie kann aber leicht durch das gerundete, ungezähnte Dorsellum von den hier 
behandelten Arten unterschieden werden. 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 365 


Clypeus stark vorgezogen (Abb. 3a); Flagellum schlank, Glieder 2-4 
beim ® länger als breit (Abb. 3b), beim d quadratisch oder wenig 
länger als breit (Abb. 3c). Mesoscutum im Zentrum mit 2 gelben 
Flecken (Abb. 3d), manchmal diese sich gegenseitig berührend, beim 
Gro hte hei dey eee ENTER END. OR ARE L. gigas 
Clypeus mässig vorgezogen (Abb. 3a’); Flagellum gedrungener, Glieder 
2-4 beim 2 wenig kürzer als breit oder quadratisch (Abb. 3b'), beim 4 
deutlich kürzer als breit (Abb. 3c'). Mesoscutum im Zentrum mit nur | 
selben Bleck (Abb Sd) bem oitiehlendee Pan aie ae L. intermedia 


4) - 


HANNES BAUR & FELIX AMIET 


Pronotum mit 2 Querkielen: einer im hinteren Viertel, der andere fast 
unmittelbar entlang des Hinterrandes (Abb. 4a). 2: Ovipositor erreicht 
nicht das 1. Gastraltergum, dieses ohne Längsfurche und sein gelbes 
Querband in der Mitte stark eingeschnürt (Abb. 4b); Längsfurche des 5. 
Gastraltergums tief, Ovipositorscheide in Ruhelage teilweise verdeckt 
CADDRAC) Entei EEE DEE EE L. biguetina 
Pronotum mit 3 Querkielen: ein kurzer ungefähr in der Mitte, gefolgt 
von zwei längeren im hinteren Viertel respektive fast unmittelbar ent- 
lang des Hinterrandes (Abb. 4a'). 9: Ovipositor erreicht das 1. Gastral- 
tergum, dieses mit Langsfurche und sein gelbes Querband in der Mitte 
nicht eingeschniirt (Abb. 4b'); Langsfurche des 5. Gastraltergums ober- 
flachlich, Ovipositorscheide in Ruhelage immer ganz sichtbar (Abb. 
ACER RCE Ss RL ee ee RAI N RR REI 0000000 5 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 367 


5)— Clypeus deutlich vorgezogen, lateral mit aufgebogenen Lappen, welche 

nach unten leicht konvergieren (Abb. 5a). 9: Längsfurche des 1. 
Gastraltergums an der Basis kaum verjüngt (Abb. 5b)......... L. dorsigera 

— Clypeus kaum vorgezogen, mehr oder weniger verrundet und lateral 

nur mit fein angedeuteten Kielen (Abb. 5a'). 9: Längsfurche des 1. 


Gastraltergums an der Basis deutlich verjüngt (Abb. 5b')....... L. bifasciata 
1! [orme ne =] 
0 5a 
rer] 
i 
en 5b 5b' 
Leucospis gigas Fabricius, 1793 Abb. la-Id, 2a', 3a-3d, 6-8, 12-14. Tabl. 


Untersuchtes Material [Schweiz: alphabetisch geordnet nach Kanton, Ort; übrige 
Länder: alphabetisch geordnet nach Land, Provinz bzw. Region, Ort]: SCHWEIZ: ohne 
Fundortsangabe 2 ? (MNHU); «Cornet»: 1 ©, 1867 (MHNG). BE: Bern: 3 2, leg./coll. M. 
Perty (NMBE). GE: Genève: 3 ©, leg. Bües (ZSM); 1 ©, 15.6.1889, 2 2, 20.7.1889, 5 9, 
20.7.1889, leg./coll. F. Chevrier (MHNG). Genève, Champel: 1 9, [?18...], 1 do, 7.[?18...], 1 
99187.018.),1°2,227.048°2] (MENG). GR? Brusio: 1° 99317.3:.8.1933 Je gIc0ll aA 
Nadig (ETHZ). VD: Chamblon, Champ Muraz: 1 2, 1942, ex Megachile parietina, leg. A. 
Sermet (ASE). Nyon: 3 ©, leg./coll. F. Chevrier (MHNG). VS: 1 2 (MSNV); 1 ®, coll. 
Naumberg (DEI); 1 9, leg./coll. FE. Frey-Gessner (NMSO); 114 ®, leg./coll. M. Paul (NMW); 
1 ©, leg./coll. O. Schmiedeknecht (NMBE); 1 ®, leg./coll. H. Tournier (MHNG); 6 2, leg. T. 
Steck (NMBE); 1 ®, leg. A. Wullschlegel (NMBE). Brig: 2 ®, 7.1914, leg./coll. A. von 
Schulthess (ETHZ). Euseigne: 9 2, 8.7.1908, 2 ©, 5.7.1911, leg. T. Steck (NMBE); 1 ©, 
87.1908, 1S 2561924 <1 2.,16:8:1929,172157.1936, 1es/collÜT. Steck (NMBA) 339 27 
31.7.1939, 1 ©, 23.7.-2.8.1942, leg./coll. A. Nadig (ETHZ); 1 ?, 2.8.1940, leg./coll. J. de 
Beaumont (MZL). Finges: 1 2, 25.7.1946, leg./coll. J. Aubert (MZL); Fully, Les Follatéres: 1 
2, 5.7.1947, leg./coll. J. Aubert (MZL). Gampel, Jeizinen: 1 9, 8.8.1998, an Sedum album, leg. 
H. Baur (NMBE). Martigny: 1 9, 7.7.1935, leg. P. Bovey (MZL); 1 2, 18.7.1936, 1 9 
25.6.1948, 1 2, 4.7.1948, 1 9, 28.6.1956, leg./coll. J. de Beaumont (MZL). Niouc: 1 9, 
17.7.1900, leg./coll. E. Frey-Gessner (MHNG). Pfynwald: 1 9, 18.6.1921, 1 2, 2.7.1921, leg. 
122 Steck (NMBE)FE 27218:641921721725,3:741922 127261924 Ona 702 SMS 
20.8.1925, 1 2, 21.7.1927, 2 2, 23.7.1927, leg./coll. T. Steck (NMBA); 6 9, Ende 7.1939, 1 2, 


368 HANNES BAUR & FELIX AMIET 


13.8.1941, 9 2, 23.7.-2.8.1942, leg./coll. A. Nadig (ETHZ). Salgesch: 1 ©, 11.7.1885, 
leg./coll. M. Paul (NMW). Savièse: 1 2, 7.-8.1906, leg. J. Jullien, coll. V. Delucchi (ETHZ); 
10 2, 7.1906, leg./coll. J. Jullien (MHNG). Sierre: 3 9, coll. A. von Schulthess (ETHZ); 1 ©, 
leg./coll. Cerutti (MZL); 1 9, leg./coll. O. Schmiedeknecht (DEI); 3 2 (MHNG); 1 ©, 6.7. 
(MHNG); 3 ©, 25.6.1880, leg./coll. E. Frey-Gessner (ZSM); 1 9, 9.7.1884, 6 2, 7.1886, leg. 
T. Steck (NMBE); 2 2, 10.7.1886, leg./coll. R. Meyer-Dür (NMBE); 2 ©, 26.2.1901, coll. A. 
von Schulthess (ETHZ); 2 2, 8.1965, leg. I. Wall (IW [nicht untersucht]). Sion: 1 9, leg./coll. 
E. Frey-Gessner (MHNG); 3 ®, 25.6.1901, coll. A. von Schulthess (ETHZ); 1 2, 1906, leg. J. 
Jullien (MHNG); 1 9, 13.6.1906, leg. Suereuse (MHNG); 1 ®, 1909, leg./coll. B. Jacob 
(MZL); 1 2, 25.6.1912, leg./coll. A. von Schulthess (ETHZ); 3 ©, 20.6.1922, 1 9, 28.6.1924, 
leg./coll. T. Steck (NMBA). St. Niklaus: 1 2, 15.7.1925, 2 2, 16.7.1925, 1 2, 19.7.1927, 
leg./coll. E. Däniker (NMBE); 1 2, 19.7.1927, leg./coll. E. Däniker (WA). Stalden: 1 9, 
26.0.1909, leg: T. Steck (NMBE); 1 2, 11.7.1923,.2 9) 14.7.1923 Ales/colSEFDaniker 
(NMBE); 1 2, 4.7.1924, leg./coll. T. Steck (NMBA). Susten: 2 9, 24.7.1886, leg./coll. M. Paul 
(NMW); 3 2, 13.7.1899, 1 2, 13.7.1899, leg./coll. B. Jacob (MZL); 1 9, 1.7.1900, leg./coll. E. 
Frey-Gessner (MHNG); Törbel: 1 2, 8.7.1924, leg. G. C. Krüger (ETHZ); Vex: 1 2, 5.7.1928, 
leg./coll. T. Steck (NMBA); Zeneggen: 1 2, 15.-29.7.1943, leg./coll. J. de Beaumont (MZL). 
ZH: Zürich: 1 ©, coll. A. von Schulthess (ETHZ); 1 9, «Chalicodoma muraria Nest», leg./coll. 
A. von Schulthess (ETHZ); 1 2, 9.1871, leg. F. Ris, coll. A. von Schulthess (ETHZ). 

FRANKREICH: Basses Alpes, Digne 1 © (ZSM); Basses Alpes, Valensole 2 2 (NMBE), 
2 2,1 d (FA); Bouches-du-Rhône, Fontvielle, Alpille's 1 © (MH); Provence, Callian 1 9 
(NMBA), 2 2 (NMBE); Provence, St-Jean 1 2 (MH). GRIECHENLAND: Hellas, Alt Korinth 1 2 
(WA); Hellas, Antikes Samiko 1 2 (WA); Hellas, Epid. Limera 1 2 (WA); Hellas, Kalogria, 
Achaia 1 2 (WA); Hellas, Leptokoria 2 2 (FB); Hellas, Mavromati, Ithome 1 & (WA); Hellas, 
Olympia, Alfios-Tal 1 9, 1 d (WA); Peloponnes, Neapoli, Kap Malea 1 2 (WA). ITALIEN: 
Aostatal, Nus 1 2, 1 4 (ES): Aostatal, Pondel 1 9 (CS): Aostatal, St. Pierre 2 2 (CS); Bozen 
2 2 (NMW); Bozen, St. Pauls 1 2 (NMW); Sizilien, Passomartino 1 2 (NMBE); Triest 1 2 
(ZSM); 1 2 (NMW); Triest, Conconello (Umg.) 4 2 (NMW); Triest, S. Primus 1 2 (NMW); 
Vinschgau, Meran 1 2 (DEI). KROATIEN: Dalmatien, Sibenik (Adriaküste) 6 2, ex Megachile 
hungarica (GR); Dalmatien, Sibenik 5 9, ex Megachile parietina (GR); Krk 1 2 (ZSM); Krk 1 
36 (ETHZ); Krk, Baska 4 9 (ZSM); Krk, Hvar 1 2 (ZSM); Krk, J. Hvar Jelsa 10 ©, 4 à 
(ZSM); Krk, Malinska 5 © (ZSM); Rijeka 2 © (ETHZ). MALTA: 3 2 (NMBE). ÖSTERREICH: 
Tirol, Klobenstein 1 2 (NMW); Tirol, «Larche» 1 2 (NMW). TUNESIEN: Sbeitla 1 2 (NMBE). 
TÜRKEI: Kayseri, Göreme 1 9 (CS). 


Diagnose ®: Körper schwarz mit folgenden Teilen gelb [bei manchen 
südeuropäischen und nordafrikanischen Individuen leuchtend orangerot]: Gesicht 
beiderseits der Scroben mit je einem Fleck (Abb. 3a); Scapus ausser an Basıs und 
Apex, manchmal auch dorsal dunkel: Flagellumglieder 1-3 an der Basis häufig leicht 
rötlich; Pronotum vorne mit langem, hinten mit etwas kürzerem Querband, oder das 
Sklerit vollständig eingerahmt; Mesoscutum mit 2 Flecken zentral und 2 länglichen 
Flecken lateral (Abb. 3d); Scutellum am Hinterrand mit in der Mitte stark einge- 
schnürtem Querband; Mesepisternum selten mit einem Fleck; Metapleuron voll- 
ständig hell; Metacoxa dorsal mit Basalfleck, ventral gelegentlich mit Apikalfleck; 
Pro- und Mesofemur apikal zu 2/3 bis 1/4; Aussenseite des Metafemurs, ausser einem 
dunklen Fleck und den Zähnen (Abb. 2a’): Tibien und Tarsen, nur Metatibia ventral 
dunkel; Gastralterga 1, 4 und 5 mit breiten Querbändern, welche in der Mitte unter- 
brochen sind; Tergum 7 lateral mit länglichen Flecken. 

Clypeus stark vorgezogen (Abb. 3a); Flagellum schlank, leicht keulenförmig, 
Glieder 2-4 länger als breit (Abb. 3b); Pronotum mit 2 Querkielen: einer im hinteren 
Viertel, der andere fast unmittelbar entlang des Hinterrandes (ähnlich L. biguetina, 
Abb. 4a): Dorsellum am Hinterrand mässig ausgerandet, die beiden Zähne mehr oder 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 369 


weniger deutlich; basaler Zahn des Metafemurs kürzer als folgende Zähne (Abb. 2a’); 
Längsfurche des |. Gastraltergums an der Basis leicht verjüngt; Längsfurche des 5. 
Tergums oberflächlich, Ovipositorscheide in Ruhelage immer ganz sichtbar (Abb. 
1d); Ovipositorscheide 0.95-1.46 mal so lang wie der Metafemur (Abb. 12), reicht bis 
ungefähr zur Hälfte, zum basalen Viertel oder zur Basis des 1. Gastraltergums (vgl. 
Variabilität 9). 

Körperlänge 7.0-17.2 mm (Abb. 13), Kopfbreite 2.15-3.95 mm (Abb. 14). 


Diagnose 4G: Gelbe Zeichnungselemente oft reduziert, die Flecken im 
Gesicht und im Zentrum des Mesoscutums oft fehlend; Querbänder auf dem Gaster 
durchgehend. 

Flagellum etwas gedrungener, Glieder 2-4 quadratisch oder wenig länger als 
breit (Abb. 3c). Form des Gasters ähnlich L. dorsigera (Abb. 2b’). 

Körperlänge 11.2-14.5 mm (Abb. 13), Kopfbreite 2.85-3.40 mm (Abb. 14). 

Variabilität 9: Aufgrund des Verhältnisses Ovipositorscheide- zu 
Metafemurlänge (O/M) können drei Formen unterschieden werden (Abb. 12), welche 
zusätzlich bis zu einem gewissen Grad in der Zeichnung und der Grösse (Abb. 13, 14) 
variieren. Die Formen 'A' und 'B' mit kurzer (O/M = 0.95-1.18, n = 79) respektive 
langer (O/M = 1.30-1.46, n = 18) Ovipositorscheide treten häufig zusammmen in 
mittel- und südeuropäischen Populationen auf. Im Wallis kommt hingegen nur eine 
dritte Form vor, welche in der Länge der Ovipositorscheide (O/M = 1.16-1.30, n = 
123) eine intermediäre Stellung einnimmt. Die Unterschiede der Walliser Population 
zu den beiden anderen Formen sind hoch signifikant (ANOVA mit Dunnett T3-Test, 
F = 992.887, n = 220, p < 0.001). Die Formen 'A' und 'B' wurden schon relativ früh 
erkannt und als separate Arten unter eigenen Namen beschrieben (Klug, 1814). Nach 
Boucek (1974) sind diese aber eindeutig als Synonyme von L. gigas zu werten. 

Verbreitung: Paläarktisch, östlich bis China (Bouéek, 1974); in 
Mitteleuropa aus allen Ländern nachgewiesen (vgl. Boucek, 1959; Madl, 1989; 
Schletterer, 1890). In der Schweiz wurde die Art ausser im Mittelland bei Bern und 
Zürich und am Jurasüdfuss bei Yverdon vor allem im Süden (Genferseebecken, GR, 
VS) gefunden (Abb. 6). Nach 1950 wurde L. gigas nur noch sehr selten (0.4% der 
Daten, 16% der Fundpunkte auf der Verbeitungskarte) und ausschliesslich im Wallis 
gesammelt. Die Verbreitung stimmt recht gut mit derjenigen der potentiellen Wirte 
überein, wobei diese in der Schweiz weiter verbreitet sind (Nordschweiz, GE, TI, GR) 
(Abb. 6). 

Höhenverbreitung: von 375 bis 1505 mii. M. (Abb. 6). 

Gefährdung: in der Nordschweiz wahrscheinlich ausgestorben, in der 
Südschweiz gefährdet und in der ganzen Schweiz stark gefährdet. Von den poten- 
tiellen Wirtsarten wird Megachile parietina (Geoffroy) in der Roten Liste (Amiet, 
1994) für die Nordschweiz als vom Aussterben bedroht, für die Südschweiz und für 
die ganze Schweiz als gefährdet eingestuft. M. pyrenaica Lepeletier, welche vor allem 
in den Alpen, selten im Jura und im Mittelland vorkommt, gilt als nicht gefährdet. Die 
Beurteilung der Gefährdung von M. parietina trifft ebenfalls recht gut auf L. gigas zu. 
Allerdings ist die Art in der Nordschweiz, wo sie nach Sulzer (1776) und Fuessly 
(1783) z. B. im Kanton Zürich mehr oder weniger regelmässig auftrat, wohl überall 
ausgestorben. Zudem ist sie im Wallis im Rhonetal stark zurückgegangen. Aufgrund 


370 HANNES BAUR & FELIX AMIET 


- Höhenverbreitung n=154 Phänologie n=90 
2000 4 
1500 
1000 
500 
— = : : 
0% 25% 50 % 75% Mai Juni Juli August September 


ABB. 6, L. gigas: Höhenverbreitung, Phänologie und Verbreitung in der Schweiz; @ Funde nach 
1950, O Funde vor 1951, potentielle Wirte (Auswertung von 100 Datensätzen von Megachile 
parietina und M. pyrenaica). 


der Daten war die Art dort früher wahrscheinlich verbreitet und relativ häufig. Gegen- 
wärtig ist sie aber nur noch von einer einzigen Stelle bei Jeizinen bekannt. Vor diesem 
Hintergrund muss L. gigas für die gesamte Schweiz als stark gefährdet eingestuft 
werden. 

Biologie: Zu den Wirten werden die folgenden Angaben gemacht, 
welche als gesichert gelten können: Megachile parietina (Amiet, Baur, Forster und 
Zettel, pers. Beob. an Nestern im Wallis; Fabre, 1886 sub «Chalicodome des 
murailles» oder «Chalicodome des galets»; Bouëek, 1970, Fahringer, 1922, Giraud & 
Laboulbène, 1877 und Schletterer, 1890 sub Chalicodoma muraria); M. pyrenaica 
(Fabre, 1886 sub «Chalicodome des hangars»; Fahringer, 1922 sub Chalicodoma), M. 
sicula (Rossi) (Schletterer, 1890 sub Chalicodoma; Bürgis, 1995). Ferner wurden die 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 371 


ABB. 7: Weibchen von L. gigas schlüpft aus einem Nest von Megachile parietina von Sibenik 
(Kroatien). Daneben die Exuvie und die Imago der ebenfalls parasitischen Anthrax ?binotatus 
Wiedemann (Diptera: Bombyliidae) (Foto G. Reder). 


folgenden Exemplare mit zuverlässigen Wirtsangaben untersucht: FRANKREICH: Kor- 
sika, Calvi (Umgebung), Stareso, 24.9.1991, ex M. sicula 1 ®, e. p. 11.-13.6.1993, 
1 9, e. p. 1994, leg. H. Bürgis (HB). KROATIEN: Dalmatien, Sibenik (Adriaküste) 6 
2, e. p. 14.-22.5.1998, ex M. hungarica (Mocsäry), 5 2, e. p. 22.-29.5.1998, ex M. 
parietina, leg. G. Reder (GR). SCHWEIZ: VD, Chamblon, Champ Muraz 1 2, 1942, ex 
M. parietina, leg. A. Sermet (ASE). ZH, Zürich 1 9, ex M. parietina, leg./coll. A. von 
Schulthess (ETHZ). 

Die Nennung weiterer Wirte von L. gigas geht zumeist auf eine einzige, oft 
recht alte Quelle zurück. So erwähnt Fahringer (1922) den Schlupf von mehreren 
Osmia bicornis (Linnaeus) und einem Exemplar von L. gigas aus einem verlassenen 
Nest von M. parietina. Da L. gigas bis zu drei Jahren überliegen kann (Bürgis, 1995), 
ist nicht auszuschliessen, dass die Entwicklung zuvor an M. parietina, und nicht an 
O. bicornis, stattfand. Mann fand «in Corsica einen tiber nussgrossen, kugeligen Bau, 
aus Thon und Sandkörnern hergestellt und an einem Zweige von Cystus salviaefolius 
befestigt» woraus Odynerus trifasciatus Fabricius (Vespidae), Osmia ‘coerulea’ 
[= ?0. caerulescens (Linnaeus)] und L. gigas schlüpften (Schletterer, 1890: 11). Hier 


372 HANNES BAUR & FELIX AMIET 


ABB. 8: Ausschnitt aus dem Habitat von L. gigas bei Jeizinen im Wallis (Schweiz) (Foto B. 
Baur). 


Osmia als Wirt anzunehmen, ist zwar naheliegend, aber möglicherweise irreführend. 
So passt die Beschreibung des «Bau» vorzüglich auf das Nest von M. sicula, ın 
diesem Fall der wahrscheinlichere Wirt. Der Parasitoid könnte M. sicula zu einem 
frühen Zeitpunkt attackiert haben, und dann, nach Jahren des überliegens, gleichzeitig 
mit der O. caerulescens geschlüpft sein, welche in der Zwischenzeit die verwaisten 
Zellen für sich benutzte. Die Wirtsangabe Anthophora garrula (Rossi) (Fahringer, 
1922 sub Podalirius) bedarf der Bestätigung, diejenige von Vespula vulgaris (Lin- 
naeus) (Rondani, 1873) ist äusserst zweifelhaft. 

Brutparasiten (Coelioxys, Stelis) werden gelegentlich als Wirte von Leucospis 
erwähnt, konnten aber bisher nicht schlüssig bestätigt werden. Boucek (1974) 
berichtet lediglich von einem Exemplar von L. gigas, das in Frankreich zusammen 
mit Coelioxys quadridentata (Linnaeus) [= C. conica (Linnaeus)] gezogen wurde. 
Amiet (pers. Beob.) beobachtete jedoch im Juli in Sambiase, Kalabrien, Italien, wie 
ein Exemplar von L. gigas mehrmals immer wieder an der gleichen Stelle das Nest 
einer Megachile sp. (parietina oder sicula) anstach. Die Untersuchung des Nestes 
ergab vier Zellen, eine verschimmelte, zwei besetzt mit mehreren Larven von ?Stelis 
nasuta (Latreille) und eine mit einer fertig entwickelten Megachile-Larve. Die 
Leucospis setzte die Spitze des Ovipositors genau über der letztgenannten Zelle an. 

Fabre (1886) schildert in seinem Aufsatz über die Leucospis eingehend das 
Verhalten von L. gigas. Eine kurze Zusammenfassung über das Eiablageverhalten 
sowie die Ontogenie einschliesslich Abbildungen des Eis und der Larvenstadien 
findet sich in Clausen (1940). Bürgis (1995) berichtet, dass L. gigas bis zu mehreren 
Jahren im Nest von Megachile sicula ausharren könne. Entgegen der verbreiteten 
Annahme (z. B. in Jacobs & Renner, 1998), wonach die fertig entwickelte Wespe auf 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 373 


die Hilfe eines zuvor schlüpfenden Wirtes, welcher einen Ausgang freimache, ange- 
wiesen sei, befreiten sich alle Exemplare selbständig aus dem Nest (Bürgis, 1995). 
Ferner konnte Reder (pers. Beob.) mehrere Exemplare von L. gigas beim Schlupf aus 
Nestern von M. parietina (Abb. 7) bzw. M. hungarica beobachten. Auch in diesem 
Fall haben alle Parasitoide mit Hilfe der Mandibeln scheinbar mühelos das Nest 
selbständig verlassen. 

Imagines erscheinen von Juni bis August (Abb. 6) und werden meist an 
xerothermen Standorten wie Steppen- und Trockenrasen (Abb. 8) angetroffen, wo sie 
in unruhigem, raschem Flug das Gelände nach Nestern der Wirte absuchen. Einzelne 
Exemplare konnten auf Blüten von Allium sphaerocephalon Linnaeus (Liliaceae) 
(Forster, pers. Beob.) und Sedum album Linnaeus (Crassulaceae) (vgl. Material) 
beobachtet werden. Nach Ansicht verschiedener Autoren (Berland, 1934a; Bytinski- 
Salz, 1963) vermehrt sich L. gigas in Europa vor allem parthenogenetisch (thelytok), 
da hier Männchen nur selten gefunden werden. Dies scheint auch für die Populationen 
in der Schweiz zuzutreffen, denn unter den insgesamt 281 untersuchten Exemplaren 
befindet sich nur ein einziges Männchen aus der Nähe von Genf. 


Leucospis intermedia Niger, 1807 Abb. 3a'-3d', 9, 12-14 


Untersuchtes Material: SCHWEIZ: GR: Brusio: 2 d, 31.7.-5.8.1935, leg./coll. A. Nadig 
(ETHZ). Grono: 3 ©, 7.1887, leg. T. Steck (NMBE); 1 2, 21.8.1934 (NMBA). Roveredo: 1 d, 
20.7.1928, leg./coll. A. Nadig (ETHZ); 1 2, 7.8.1935, leg./coll. T. Steck (NMBA). VD: 
Buchillon: 1 ©, 17.7.1930, leg. P. Bovey (ETHZ). Nyon: 1 6, leg./coll. F. Chevrier (MHNG). 
VS: ohne Fundortsangabe 1 9, leg. A. Wullschlegel (NMBE); 1 d, leg./coll. E. Frey-Gessner 
(NMW); 1 9, leg./coll. H. Tournier (MHNG). Aven: 1 @, 28.5.1992, leg./coll. F. Amiet 
(NMSO). Branson: 2 4, 30.5. (MHNG); 1 d, 23.6.1964, 1 ?, 29.7.1964, leg./coll. J. de 
Beaumont (MZL). Euseigne: 1 6, 5.7.1911, leg. T. Steck (NMBE); 2 2, 31.7.1939, leg./coll. 
A. Nadig (ETHZ). Evolène: 1 4, 14.7.1911, leg. T. Steck (NMBE). Fully, Les Follatères: 1 4, 
1929 (MHNG); 2 2, 11.7.1933, 1 2, 21.7.1933, 1 8,1 2, 27.6.1942, leg. P. Bovey (MZL); 2 
221824126194 I 25 195.1948, leg/coll. A. Nadis (EIHZ); 1 2, 14.6.1951, leg./coll. J. 
Aubert (MZL); 1 2, 29.7.1964, leg./coll. J. de Beaumont (MZL). Grimentz: 1 ©, 27.7.- 
12.8.1944, leg./coll. J. de Beaumont (MZL). Val d'Hérens: 1 4, 26.6.1894, leg./coll. M. Paul 
(NMW). Martigny: 1 9, leg./coll. B. Jacob (MZL); 1 3, 1 2, 1.-4.6.1875, 1 9, 7.-8.6.1875, 
leg./coll. E. Frey-Gessner (ZSM); 1 8,5 9, 15.6.1889, leg. T. Steck (NMBE); 1 9, 5.7.1933, 
leg./coll. J. de Beaumont (MZL). Mazembroz: 1 6, 14.5.1934, leg. P. Bovey (ETHZ); 1 d, 
18:541973712.870:0:1976,, 1.8, 15:8:1977, 273, 2.619887 leg. E. Amict (EA) Nioucs 279, 
12.8.1908, leg. T. Steck (NMBE). Ried bei Morel, Riederalp: 1 2, 16.7.1921, leg./coll. E. 
Däniker (NMBE). Sierre: 1 d, 3 ©, coll. A. von Schulthess (ETHZ); 1 2, leg./coll. A. von 
Schulthess (ETHZ); 1 d, leg./coll. O. Schmiedeknecht (NMW); 1 à, leg./coll. O. Schmiede- 
knecht (MNHU); 1 4, 16.7.1880, ex Megachile sp., leg./coll. E. Frey-Gessner (MHNG); 1 4, 
9.7.1884, leg./coll. A. von Schulthess (ETHZ); 1 d, 22.7.1884, 1 9, 7.1886, leg. T. Steck 
(NMBE); 1 d, 24.6.1885, leg./coll. M. Paul (MHNG); 1 d, 27.6.1886, 3 2, 1887, leg. M. Paul, 
coll. A. von Schulthess (ETHZ). Sion: 1 2, leg./coll. F. Chevrier (MHNG); 1 6, 8.7.1884, leg. 
T. Steck (NMBE); 1 2, 9.1912 (NMBA). St. Niklaus: 1 2, 19.7.1927, leg./coll. E. Däniker 
(NMBE). Stalden: 1 6, 26.6.1909, 1 d, 1 2, 27.6.1909, 3 d,2 9, 28.6.1909, leg. T. Steck 
(NMBE). Vex: 2 d, 21.6.1935, leg./coll. T. Steck (NMBA). Zeneggen: 1 ©, 20.7.1986, leg. F. 
Amiet (FA). 

FRANKREICH: Py-Montet 1 d (MH); Tence 1 2, ex Osmia emarginata (MHNG). GRIE- 
CHENLAND: Hellas, Kallogria, Achaia 2 £ (WA); Peloponnes, Blafonisoa 1 2 (WA). ITALIEN: 
Aostatal, W Arvier 1 & (CS); Aostatal, Sarre 1 £ (CS); Aostatal, St. Pierre 1 2 (CS); Bozen 2 
2,13 (NMW); Bozen 2 9, 2 gd (ETHZ); Puglia, Mte. Gargano, NW M. S. Angelo 1 2 
(ETHZ); Sizilien, Siracusa 1 2 (NMBE); Vinschgau, Latsch 1 2 (ES). KROATIEN: Dalmatien, 


374 HANNES BAUR & FELIX AMIET 


Split 6 2 (NMW); Krk 2 ? (ETHZ); Krk, Hvar 2 ® (ZSM); Krk, J. Hvar Jelsa 1 2 (ZSM). 
ÖSTERREICH: Kärnten, Villach 1 2 (NMW); Tirol, «Larche» 1 2 (NMW). 

Diagnose ® ähnlich der von L. gigas, aber: Flagellumglieder 1-3 nie 
rötlich; Pronotum vorne mit kurzem, hinten mit langem Querband (ähnlich L. 
dorsigera, Abb. 4a'), oft das Sklerit vollständig eingerahmt; Mesoscutum mit nur 1 
Fleck zentral; Propodeum selten mit je einem kleinen Fleck lateral; Querband auf 
Gastraltergum 4 manchmal kaum unterbrochen. 

Clypeus mässig vorgezogen (Abb. 3a’); Flagellum gedrungener als bei L. 
gigas, fast zylindrisch, Glieder 2-4 wenig kürzer als breit oder quadratisch (Abb. 3b’); 
Längsfurche des |. Gastraltergums normalerweise nicht verjüngt; Ovipositorscheide 
länger, 1.46-1.93 mal so lang wie der Metafemur (Abb. 12), reicht meist mindestens 
bis zum Hinterrand des Scutellums, oft bis zu dessen Mitte. 

Körperlänge 4.5-12.3 mm (Abb. 13), Kopfbreite 1.45-3.10 mm (Abb. 14). 

Diagnose d: Gelbe Zeichnungselemente oft reduziert, der Fleck im 
Zentrum des Mesoscutums oft fehlend; Querbänder auf dem Gaster durchgehend. 

Flagellum etwas gedrungener, Glieder 2-4 deutlich kürzer als breit (Abb. 3c’). 
Form des Gasters ähnlich L. dorsigera (Abb. 2b’). 

Körperlänge 6.8-10.2 mm (Abb. 13), Kopfbreite 2.05-2.85 mm (Abb. 14). 

Verbreitung: Paläarktisch, östlich bis Tadschikistan (Bouéek, 1974); in 
Mitteleuropa aus allen Ländern nachgewiesen (vgl. Boutek, 1959; Madl, 1989, 1990; 
Schletterer, 1890). In der Schweiz ist die Art auf das Wallis, das Genferseegebiet und 
die Alpensüdtäler beschränkt (Abb. 9). Nach 1950 wurde L. intermedia nur noch 
gelegentlich (19% der Daten, 22% der Fundpunkte auf der Verbreitungskarte) und 
ausschliesslich im Wallis gesammelt. Die Verbreitung stimmt, mit Ausnahme des 
Genferseebeckens, recht gut mit derjenigen des potentiellen Wirtes überein, wobei 
dieser in der Schweiz weiter verbreitet ist (Mittelland, Alpen) (Abb. 9). 

Höhenverbreitung: von 290 bis 1925 mii. M. (Abb. 9). 

Gefährdung: nicht gefährdet. Der potentielle Wirt Osmia mustelina 
Gerstäcker hat keinen Eingang in die Rote Liste (Amiet, 1994) gefunden und gilt in 
der Schweiz als nicht gefährdet. 

Biologie: Als Wirt wird von verschiedenen Autoren bisher nur Osmia 
emarginata Lepeletier angegeben (z. B. Berland, 1934b für Frankreich; Bouéek, 1959 
für Tschechien; Giraud, 1858 für Österreich; Madl, 1990 für Deutschland). Allerdings 
werden in jüngerer Zeit die Populationen östlich von Frankreich als eigene Art, O. 
mustelina, aufgefasst. Daher kommt als Wirt von L. intermedia in der Schweiz vor 
allem diese Osmia-Art in Frage. Ferner wurden die folgenden Exemplare mit Wirts- 
angabe untersucht: FRANKREICH: Haute-Loire, Tence 1 9, 16.7.1925, «parasite d’O. 
emarginata» (MHNG); SCHWEIZ: VS, Sierre 1 d, 16.7.1880, «nid de Megachile No. 
9320, e. p. 4. 1881», leg./coll. E. Frey-Gessner (MHNG); SPANIEN: Soria, Beraton 
1 9, 22.6.1994, an Nest von O. emarginata, leg. F. Amiet (FA). Die Wirtsangabe von 
Frey-Gessner bedarf der Bestätigung! 

Imagines erscheinen von Mai bis August (Abb. 9) und werden meist an xero- 
thermen Standorten wie Steppen- und Trockenrasen angetroffen, wo sie in raschem 
Flug das Gebiet nach Nestern ihrer Wirte absuchen. Frühere Vermutungen, dass sich 
L. intermedia in Europa vor allem parthenogenetisch (thelytok) fortpflanze (Boucek, 
1964), konnten nicht bestätigt werden, da Männchen überall recht zahlreich gefunden 
wurden. 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 375 


Höhenverbreitung n=84 50%, Phanologie n=62 


25% + 


0% 


r 7 
50 % 75% Mai Juni Juli August September 
SEI je à 
D | AS Sy i> 
Gx be? ENO. A 
\ | ee: me 
> G % = / ZEN 
= 1 TER < > SN Ü x 
È LR 
7 ¢ \ ARI TAO bri E 
1 \ EN 
= ) = LE DIA I 
RA ; vi 7 
À \ ) —_ \ \ 
\ Es LY RE \ / 
N A E. Zee \ A 
SZ É = Wr JS 
O oa EG = ou A 
Le OL | fh 
ee Le - Ò 
ot | 8 Ley = 
> EEA \ 
we 9/0 5 
2) à 


ABB. 9, L. intermedia: Höhenverbreitung, Phänologie und Verbreitung in der Schweiz; @ Funde 
nach 1950, © Funde vor 1951, potentieller Wirt (Auswertung von 38 Datensätzen von Osmia 
mustelina). 


Leucospis dorsigera Fabricius, 1775 Abb. 2b’, 4a'-4c', Sa, Sb, 10, 12-14 


Untersuchtes Material: Schweiz: ohne Fundortsangabe 1 4, leg. Morawitz, coll. Gerst. 
(MNHU). BE: Batterkinden: 1 2, 25.7.1887, leg./coll. T. Steck (NMBA). Bern: 2 2, leg./coll. 
M. Perty (NMBE); 1 9, 1.8.1897, leg. T. Steck (NMBE). Bern, Kirchenfeld: 1 3, 9.6.1922, 
leg./coll. T. Steck (NMBA). Biel: 1 2, 29.7.1888, leg./coll. T. Steck (NMBA). Lyss: 1 2, 
26.6.1908, leg. T. Steck (NMBE); 1 ®, 26.8.1923, leg./coll. T. Steck (NMBA). La Neuveville: 
1 2, 13.7.1897, leg./coll. B. Jacob (MZL). GE: l'Allondon: 1 2, 29.6.1957, 2 2, 20.6.1959, 
leg./coll. Steffen (MHNG). Genève, Champel: 1 2, 17.9.[?18...] (MHNG). Genève, Corsier: 1 
2, 14.7.1985, leg./coll. Steffen (MHNG). Russin: 1 ©, 6.8.1995, leg. F. Amiet (FA). Satigny, 
Peney: 1 9, 30.5.1875 (MHNG); 1 6, 21.6.1875 (MHNG); 1 ©, 7.1875 (MHNG); 1 28, 
1.7.1875 (MHNG). GR: Grono: 4 2, 7.1885, leg. T. Steck (NMBE); 1 9, 27.6.1921, 2 ®, 
30.6.1923, 1 2, 19.8.1924, leg./coll. T. Steck (NMBA). Klosters: 1 £ (ETHZ). Landquart: 1 
3, 8.6.1996, 1 2, 14.6.1996, an Osmia adunca Brutplatz, leg. H. Tinner-Guler (HT). Mesocco: 
1 2, 14.7.1927, leg./coll. T. Steck (NMBA). Roveredo: 1 2, 19.8.1924, 1 9, 18.-20.6.1926, 


376 HANNES BAUR & FELIX AMIET 


1 2720741928,1 8,2 2, 23.7.-3.8.1933, 1 2, 17.8.1949 IegJeoll. Ay Nadig (ERHZ) MINCE 
1 2, 18.8.1934, 1 gd, 5.8.1935, 1 2, 7.8.1935, leg./coll. T. Steck (NMBA). Roveredo, S. 
Vittore: 1 2, 3.-10.8.1942, leg./coll. A. Nadig (ETHZ). NE: Auvernier: 1 ©, 7.9.1907, leg./ 
coll. B. Jacob (MZL). SH: Merishausen: 1 9, 9.8.1999, leg./coll. G. Bächli (NMBC). SO: 
Dornach: 1 9, 3.8.1995, leg./coll. F. Amiet (FA). Solothurn: 1 ©, 4.8.1995, leg./coll. F. Amiet 
(FA). TI: Acquarossa: 1 2, 9.-10.1951, leg./coll. Lautner (NMBA). Agno: 2 6, 1 9, 
15.7.1944, leg./coll. J. de Beaumont (MZL). Bignasco: 1 2, 1905, leg. T. Steck (NMBE). 
Capolago: 1 9, 12.7.1944, leg./coll. J. de Beaumont (MZL). Chiasso, Pedrinate: 1 9, 
16.8.1997, leg. F. Amiet (FA); Claro: 1 ®, 6.8.1996, leg. A. Salvioni (AS). Gordola: 1 à, 
26.7.1933, leg./coll. A. Nadig (ETHZ). Losone: 1 2, 30.6.-2.7.1959, leg./coll. J. de Beaumont 
(MZL). Maroggia: 1 6, 8.[719...], 1 2, 27.8.[719...], 1 2, 8.1919, leg. G. C. Krüger, coll. A. 
von Schulthess (ETHZ). Minusio: 1 d, 4 ®, 30.6.1953, 1 ©, 30.6.1953, an Apiaceae, 2 9, 
7.7.1953, leg./coll. A. Nadig (ETHZ). Novaggio: 1 2, 6.1911, coll. A. von Schulthess (ETHZ); 
1 d, 26.6.1920, leg./coll. T. Steck (NMBA). VD: Bonvillars, 1 9, 3.8.1981, leg./coll. Steffen 
(MHNG). Commugny: 1 ®, 24.7.1949, 1 2, 1.8.1954, 1 2, 21.7.1948, 1 2, 6.8.1950, 1 2, 
19.7.1949, 1 9, «Eté» 1946, 1 ?, 24.7.1955, leg./coll. Steffen (MHNG). Cudrefin: 1 9, 
15.8.1926, leg./coll. T. Steck (NMBA). La Sauge: 1 ?, 10.8.1959, leg./coll. J. de Beaumont 
(MZL). VS: 1 à, coll. A. von Schulthess (ETHZ): 1 d, coll. H. Tournier (MHNG). Chandolin: 
1 2, 5.7.1968 (MHNG); 1 9, 7.7.1968 (MHNG). Euseigne: 1 2, 3.7.1897, 1 2, 6.8.1933, leg. 
T. Steck (NMBE); 1 9, 27.6.1924, 1 9, 6.8.1932, 1 2, 8.8.1932, leg./coll. T. Steck (NMBA); 
1 3, 25.-26.7.1939, 1 gd, 12.6.1948, leg./coll. A. Nadig (ETHZ). Fully, Les Follatéres: 1 à, 
23.7.-2.8.1942, leg./coll. A. Nadig (ETHZ). Hohtenn: 1 ®, 29.9.1987, 1 3, 15.9.1991, leg. F. 
Amiet (FA); 1 ©, 26.7.1994, leg. S. Ungricht (ETHZ). Leuk, Platten, Rotten: 1 2, 10.8.1997, 
leg. B. Merz (ETHZ). Martigny: 1 6, 15.6.1889, 1 d, 15.6.1889, leg. T. Steck (NMBE); 1 6, 
18.7.1936, 1 ©, 29.6.1946, leg./coll. J. de Beaumont (MZL). Mazembroz: 1 ®, 6.6.1976, leg. 
F. Amiet (FA). Morel: 1 ©, 1.8.1939, leg./coll. A. Nadig (ETHZ). Niouc: 1 d, 12.7.1908, leg. 
T. Steck (NMBE). Pfynwald: 1 d, 28.8.1885, leg. M. Paul, coll. A. von Schulthess (ETHZ); 1 
2, 9.7.1975, leg. F. Amiet (FA); 1 2, 15.7.1977, leg. F. Amiet (FA); 1 ©, 1989, leg. M. Paul, 
coll. A. von Schulthess (ETHZ); 3 2, 13.8.1997, leg. B. Merz (ETHZ). Sierre: 2 9, coll. A. 
von Schulthess (ETHZ); 1 à, 7.7.[718...], 1 ©, 1.8.[218...] (MHNG); 1 ©, 20.7.1884, les. T. 
Steck (NMBE); 3 ®, 1887, 2 9, 7.9.1887, leg. M. Paul, coll. A. von Schulthess (ETHZ). 
Sierre, Le Glarey: 1 9, 12.7.1905, leg. J. Jullien (MHNG). Sion: 1 2, 25.6.1912, leg./coll. A. 
von Schulthess (ETHZ). St. Niklaus: 1 ©, 15.7.1925, leg./coll. E. Däniker (NMBE). Susten: 1 
2, leg./coll. B. Jacob (MZL); 1 ®, 7.8.1887, leg. M. Paul, coll. A. von Schulthess (ETHZ). ZH: 
Zürich; 1 2, coll. A. von Schulthess (ETHZ); 1 2, 9.1870, leg. F. Ris, coll. A. von Schulthess 
(ETHZ). Zürich, Albisgiietli: 1 9, 31.7.1995, an Daucus carota, leg. S. Ungricht (ETHZ). 

DEUTSCHLAND: Baden-Wiirttemberg, Freiburg i. B. 1 2, 2 d (ETHZ); Baden-Wiirttem- 
berg, Kaiserstuhl 4 2, 1 & (ETHZ); Bayern, Dudenhofen 1 & (MH); Bayern, Eichstätt 2 9 
(ZSM); Bayern, Nürnberg, Laufamholz 1 2,2 4 (ZSM); Hessen, Flörsheim-Dalsheim 1 9 
(GR); Landau 1 © (ZSM); Thüringen, Frankenhausen, Kyffhäusern 2 © (DEI). FRANKREICH: 
Basses Alpes, Valensole 1 £ (NMBE); Bouches-du-Rhône, St-Martin de Crau 1 2, 1 4 (MH); 
Gard, Nimes 1 2 (ZSM); Provence, Callian 1 2 (NMBE); Roussillon, Argelès-sur-Mer 1 2, 
1 3 (FA). ITALIEN: Abruzzi, L'Aquita, Fonte Cerreto 1 d (ZSM); Aostatal, Sarre 1 9 (CS); 
Bozen 1 d (NMBA), 1 2 (DEI), 10 2,8 d (NMW); Bozen, St. Pauls 23 2 (NMW); Lavagna 
1 2 (ZSM); Lecco, Mte. Baro 1 9 (ETHZ); Lecco, Pusiano 12 2 (ETHZ); Sizilien, Siracusa 
1 2 (NMBE); Triest, Conconello 2 2 (NMW); Vinschgau, Meran 1 9 (ZSM). KROATIEN: Krk, 
Baska 2 2, 2 6 (ZSM); Krk, Hvar 3 2, 3 6 (ZSM); Krk, Rudina 2 2 (ZSM). ÖSTERREICH: 
Burgenland, Winden 29 9, 5 4 (NMW); Kärnten, Podcetrtek 1 2 (ZSM); Niederösterreich, 
Furth 1 ¢ (NMW); Oberösterreich, Linz 1 d (NMW); Salzburg, Glau-Moos am Heustadl 1 9, 
3 d (ZSM); Salzburg, Porsch 1 2 (ZSM); Wien, Mauer 2 & (NMW). SLOWENIEN: Ljubljana 
[Laibach, Carniolia] 3 2,1 & (ZSM). 


Diagnose 9: Körper schwarz mit folgenden Teilen gelb: Scapus von 
dunkel bis ganz hell; Pronotum vorne mit kurzem, hinten mit langem Querband (Abb. 
4a‘), manchmal das Sklerit vollständig eingerahmt; Mesoscutum selten mit 2 läng- 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 377 


lichen Flecken lateral; Scutellum am Hinterrand mit schmalem bis mässig breitem 
Querband; Metapleuron teilweise bis vollständig hell; Pro- und Mesocoxa vereinzelt 
mit kleinem Fleck, Metacoxa ventral oft mit variablem Apikalfleck; Pro- und 
Mesofemur apikal zu 1/3 bis 1/10, Metafemur meistens basal und apikal mit je einem 
Fleck, diese manchmal breit verbunden; Tibien und Tarsen grösstenteils: Protibia oft 
lateral, Metatibia immer ventral dunkel; Gastralterga 1, 4, 5 und 7 mit variabler 
Zeichnung (Abb. 4c'): Terga 1 und 5 mit breitem Querband, beim ersten durch die 
Längsfurche unterbrochen (Abb. 5b); Terga 4 und 7 mit lateralen Flecken, welche 
manchmal reduziert sind und auf Tergum 4 gelegentlich sogar fehlen. 

Clypeus deutlich vorgezogen, lateral mit aufgebogenen Lappen, welche nach 
unten leicht konvergieren (Abb. 5a); Flagellum schlank, leicht keulenförmig, Glieder 
2-4 quadratisch oder länger als breit; Pronotum mit 3 Querkielen: ein kurzer ungefähr 
in der Mitte, gefolgt von zwei längeren im hinteren Viertel respektive fast unmittelbar 
entlang des Hinterrandes (Abb. 4a’); Dorsellum am Hinterrand nur leicht ausgerandet, 
Zähne daher undeutlich; basaler Zahn des Metafemurs breit, deutlich länger als die 
folgende Reihe sehr kleiner Zähne (ähnlich L. biguetina, Abb. 2a"); Längsfurche des 
1. Gastraltergums an der Basis kaum verjüngt (Abb. 5b); Längsfurche des 5. 
Gastraltergums oberflächlich, Ovipositorscheide in Ruhelage immer ganz sichtbar 
(Abb. 4c'); Ovipositorscheide 1.53-1.92 mal so lang wie der Metafemur (Abb. 12), 
reicht normalerweise mindestens bis zur Basis des |. Tergums, oft bis zum Scutellum. 

Körperlänge 5.7-13.2 mm (Abb. 13), Kopfbreite 1.60-3.15 mm (Abb. 14). 

Diagnose d: Gelbe Zeichnungselemente mässig bis stark reduziert, 
kleine Exemplare gelegentlich bis auf helle Stellen an Pronotum, Beinen und Gaster 
vollständig dunkel; Querbänder auf dem Gaster oft durchgehend. 

Flagellum leicht gedrungener. Gaster im basalen Drittel leicht verschmälert 
(Abb. 2b’). 

Körperlänge 5.0-10.5 mm (Abb. 13), Kopfbreite 1.35-2.55 mm (Abb. 14). 

Verbreitung: Paläarktisch, östlich bis Tadschikistan (Bouëek, 1974); in 
Mitteleuropa aus allen Ländern nachgewiesen (vgl. Boucek, 1959; Madl, 1989; 
Schletterer, 1890; Schmid-Egger, 1995; Schmidt, 1969; Wolf, 1953). In der Schweiz 
kommt die Art an wenigen Stellen im Jura, Mittelland und Graubünden und an recht 
zahlreich Orten in der Südschweiz (GE, VS und TI) vor (Abb. 10). Nach 1950 wurde 
L. dorsigera nur manchmal (26% der Daten, 42% der Fundpunkte auf der Ver- 
breitungskarte) gesammelt. Die Verbreitung stimmt recht gut mit derjenigen der 
potentiellen Wirte überein, wobei diese zusammen ein deutlich grösseres Gebiet 
besiedeln (z. B. die Alpen und das Engadin) (Abb. 10). 

Höhenverbreitung: von 240 bis 1125 m i. M. (Abb. 10). 

Gefährdung: nicht gefährdet. Die meisten der potentiellen Wirte (vel. 
Biologie) werden in der Roten Liste (Amiet, 1994) als nicht gefährdet aufgeführt. 

Biologie: Aus Nestern von Anthidium diadema Latreille (Fabre, 1886), A. 
strigatum (Panzer) (Saunders, 1875 sub A. contractum), Osmia adunca (Panzer) 
(Amiet, pers. Beob.; Grandi, 1961; Müller er al., 1997; Westrich, 1989), O. bicornis 
(Linnaeus) (Boucek, 1970; Le Goff, 1999 und Schletterer, 1890 sub O. rufa), O. 
fedtschenkoi (Morawitz) (Herting, 1977), O. niveata (Fabricius) (Herting, 1977 und 
Wolf, 1953 sub O. fulviventris) und O. tricornis Fabricius (Le Goff, 1999) gezogen. 
Ferner wurde das folgende Exemplar mit zuverlässigen Wirtsangaben untersucht: 


378 HANNES BAUR & FELIX AMIET 


Höhenverbreitung n=122 50% = Phänologie n=93 
2000 | 
1500 
25% « 
1000 a 
0% — 
0% 25% 50 % 75% Mai Juni Juli August September 
= Sa Sì 
CIT En i 
= SI è —- à 2 
— Ù 1 
SE \ mn i 
7 — + 0 IN ee 
4 ele Ss MS mn 
€ = / pe È oe { N 
Va i 9 9 > “8 i E = IS NR ) 
È 7 } \ | } Y = << e =, ea 
A SI Po LC 
ie = N L = / IS oo pe 
INSEE \ A Re N 
7? RE N fer = f \ I ae F L Ne, 
Sea \ ra r > \ we 
à 4 Ex | i) ee i 
2 ES > { CE £ 
- eo N \ . ) a Do roe 
( 19) EN | O 
® 2 JIA | = 
fey À tr re 
(È ENIT AN DINE nd (OF, 
In = egli O. A O oo fo 
ir RES 5 | i my s 4 
ER OO, 
zu 


ABB. 10, L. dorsigera: Höhenverbreitung, Phänologie und Verbreitung in der Schweiz; 
@ Funde nach 1950, O Funde vor 1951, potentielle Wirte (Auswertung von 213 Datensätzen 
von Anthidium strigatum, Osmia adunca, O. bicornis, O. ligurica und O. niveata). Der Fundort 
bei Merishausen (SH) konnte auf der Verbreitungskarte nicht mehr berücksichtigt werden. 


ITALIEN: Toskana, Volterra 1 9, 10.1985, ex Nest Osmia ligurica Morawitz, leg. F. 
Amiet (FA). 

Reder (pers. Beob.) fing in seinem Garten ein Weibchen an Bruthölzern, worin 
seit vielen Jahren Osmia cornuta (Latreille), O. parietina Curtis, O. bicornis und 
Megachile ericetorum Lepeletier brüteten. Grandi (1961) beobachtete L. dorsigera an 
alten Gängen von Bostrichiden und Cerambyciden (Coleoptera), die von Bienen der 
Gattung Eriades [= Osmia] besiedelt waren. Die Angabe von Bostrichus monacha 
(Fabricius) (Coleoptera: Bostrichidae) in Herting (1973) ist gewiss ebenfalls auf die in 
den Gängen des Käfers nistende Biene zu beziehen. 

Imagines erscheinen von Mai bis September (Abb. 10) und werden meist an 
offenen, sonnigen Standorten wie Trocken- und Halbtrockenrasen, aber auch grösse- 
ren Waldlichtungen angetroffen. Einzelne Exemplare konnten auf Blüten von Daucus 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 379 


carota Linnaeus und anderen Doldengewächsen (Apiaceae) sowie Euphorbia (Eu- 
phorbiaceae) beobachtet werden (vgl. Material; Amiet, Burger, Merz, pers. Beob.; 
Chevrier, 1870; Pagliano, 1998). Aufgrund der zahlreich nachgewiesenen Männchen 
darf angenommen werden, dass sich L. dorsigera in der Schweiz in erster Linie 
bisexuell (arrhenotok) fortpflanzt. 


Leucospis biguetina Jurine, 1807 2a", 4a-4c, 11-14 


Untersuchtes Material: Lectotypus 9, ohne Fundortsangabe [Herkunft nach Boucek 
(1974): Schweiz], coll. L. Jurine (MHNG). Schweiz: VS: ohne Fundortsangabe: I 2, coll. 
Naumberg (DEI). Ausserberg: 1 £, 29.7.1933, leg. M. Naef (WL). Chandolin, Soussillon: 2 9, 
30.6.1921, leg./coll. T. Steck (NMBA). Euseigne: 1 9, 14.8.1933, leg. M. Naef (WL). 
Grengiols: 1 ©, 22.6.1998, leg./coll. F. Amiet (NMBE). Grimentz: 1 ©, 17.7.1978, leg./coll. F. 
Amiet (NMBE). Hérémence: 1 9, 23.7.1992, leg. F. Amiet (FA). Hohtenn: 2 2, 10.8.1930, 2 
2, 21.8.1932, 1 2, 29.7.1934, 2 2, 7.7.1935, leg. M. Naef (WL); 1 2, 29.7.1987, leg. F. Amiet 
(FA); 1 ®, 29.7.1987, leg./coll. F. Amiet (NMBE). Hohtenn, Ladu: 1 ©, 15.7.1991, leg. F. 
Amiet (FA). Lens: 1 9, 11.7.1967, 1 ®, 13.7.1967, 4 2, 16.7.1967, an Apiaceae (MHNG); 1 
2, 22.7.1927, leg. T. Steck (NMBE). Martisberg: 1 2, 21.7.1992, leg./coll. F. Amiet (NMBE). 
Niedergesteln: 1 9, 8.8.1998, leg. F. Amiet (FA). Niouc: 2 9, 12.7.1908, leg. T. Steck 
(NMBE). Pfynwald: 1 9, 29.6.1924, leg./coll. T. Steck (NMBA); 1 @, 18.7.1958, 1 2, 
2.8.1960, leg. Daicker (MHNG); 1 2, 7.1958, 1 ©, 10.7.1975, leg./coll. F. Amiet (NMSO); 1 
M0 7107502 77-1975 1,2 ,8:7.1979 leg; F'Amet (FA) M MO T7 197ÉMeL colle: 
Amiet (NMBE); 1 @, 6.7.1997, leg. B. Merz (NMBE); 1 2, 12.8.1997, leg. B. Merz (ETHZ). 
Sierre: 2 ©, coll. A. von Schulthess (ETHZ); 1 ©, leg./coll. O. Schmiedeknecht (DEI); 2 9, 
leg. O. Schmiedeknecht (NMW); 1 9, leg./coll. O. Schmiedeknecht (MNHU); 1 2, leg./coll. 
O. Schmiedeknecht (ETHZ); 1 2, leg./coll. T. Steck (NMBA); 1 ® (DEI); 1 2 (MHNG); 1 2, 
12.7.[?18...] (MHNG); 1 2, 15.7.1884, leg. H. Friese, coll. A. von Schulthess (ETHZ); 1 9, 
6.9.1887, leg. M. Paul, coll. A. von Schulthess (ETHZ); 1 2, 18.7.1900, 1 2, 29.6.1901, 3 ©, 
30.6.1901, leg./coll. E. Frey-Gessner (MHNG); 3 2, 26.6.1901, coll. A. von Schulthess 
(ETHZ); 2 2, 5.7.1908, leg. T. Steck (NMBE). Simplon, Berisal: 1 2, 29.7.1939, leg./coll. A. 
Nadig (ETHZ). Sion: 1 ©, 30.7.1892, leg./coll. T. Steck (NMBA); 1 2, 30.8.1892, leg. T. 
Steck (NMBE). Sion, Château de la Soie: 2 9, 9.7.1968, an Euphorbia (MHNG); 1 ©, 
13.7.1968, an Euphorbia (MHNG). St. German: 1 9, 23.7.1970, leg./coll. F. Amiet (NMBE). 
St. Niklaus: 1 2, 15.7.1925, leg./coll. E. Däniker (NMBE). Stalden: 1 2, 30.7.1924, leg./coll. 
T. Steck (NMBA); 2 2, 9.8.1933, leg. M. Naef (WL); 1 2, 7.7.1997, leg. G. Carron (GC [nicht 
untersucht]). Visp-Stalden: 1 2, 31.7.1932, 1 2, 8.8.1933, 1 2, 21.7.1935, leg. M. Naef (WL). 
Zeneggen: 1 9, 14.7.1970, 1 9, 23.7.1972, 1 2, 8.7.1973, leg. F. Amiet (FA); 1 2, 22.7.1986, 
leg./coll. F. Amiet (NMBE). 

FRANKREICH: Ariège, Verdun 1 d (SMNS); Korsika, Vizzavona 1 © (MHNG). GRIE- 
CHENLAND: Hellas, Oros Killini, Ano Trikala 1 9 (WA). ITALIEN: Postumia 1 £ (DEI); Sizilien, 
Taormina 2 d (ETHZ), 3 2,2 8 (ZSM); Vinschgau, Kastelbell 1 3 (ES); Vinschgau, Latsch 1 
dé (ES); Vinschgau, Malserheide 1 SR (EIIHZ); Vinschgau, Schluderns 3 9, 3 d (BZOL). 
KROATIEN: Krk, Baska 1 2 (ZSM). ÖSTERREICH: Burgenland, Piesting 1 £ (NMW); Bur- 
genland, Winden 10 2 (NMW); Wien, Donau-Au | £ (BZOL). SLOWENIEN: Istrien, Portoroz 
[Pirano] 1 3 (ZSM). SPANIEN: Leon, Ponferrada 1 £ (CS). TÜRKEI: Kayseri, Göreme 1 £ (CS); 
Taurus, Marasch 1 9 (ZSM). 


Diagnose ® ähnlich der von L. dorsigera, aber: Gelbe Zeichnung an den 
Beinen meist leicht rötlich verfärbt; Pronotum vorne mit langem, hinten mit kurzem 
Querband (Abb. 4a); Mesoscutum meist mit zwei Flecken im zentralen Bereich, 
durchwegs mit zwei länglichen Flecken lateral (ähnlich L. gigas, Abb. 3d); Scutellum 
am Hinterrand mit in der Mitte stark eingeschnürtem Querband; Metacoxa höchstens 
ventro-lateral mit kleinem undeutlichen Apikalfleck; Femora apikal zu 1/2 bis 1/4 


380 HANNES BAUR & FELIX AMIET 


hell, Metafemur manchmal basal mit zusätzlichem Fleck (Abb. 2a"); Tibien und 
Tarsen hell, nur die beiden ventralen Kiele der Metatibia proximal dunkel; Gastral- 
terga 1, 4, 5 und 7 mit variablen Querbändern (Abb. 4c), auf den Terga | und 4 
durchgehend, auf den beiden hinteren Terga in der Mitte unterbrochen; erstes Quer- 
band in der Mitte stark eingeschniirt (Abb. 4b). 

Clypeus kaum vorgezogen und mehr oder weniger verrundet (ahnlich L. 
bifasciata, Abb. 5a‘); Flagellum stärker keulenförmig, Glieder 2-4 wenig kürzer als 
breit oder quadratisch; Pronotum mit 2 Querkielen: einer im hinteren Viertel, der 
andere fast unmittelbar entlang des Hinterrandes (Abb. 4a); Dorsellum stärker aus- 
gerandet, mit 2 deutlichen Zähnen am Hinterrand; Gastraltergum 1 ohne Längsfurche 
(Abb. 4b), diejenige des 5. Tergums tief, so dass Ovipositorscheide in Ruhelage 
teilweise verdeckt ist (Abb. 4c); Ovipositorscheide kürzer, 1.02-1.23 mal so lang wie 
der Metafemur (Abb. 12), reicht bis zum basalen Fünftel des 5. Tergums. 


Höhenverbreitung n=64 50% = Phänologie n=69 


0% 25% 50 % 75% Mai Juni Juli August September 


ABB. 11, L. biguetina: Höhenverbreitung, Phänologie und Verbreitung in der Schweiz: @ Funde 
nach 1950, O Funde vor 1951, potentielle Wirte (Auswertung von 15 Datensätzen von Osmia 
acuticornis und O. tridentata). 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 38] 


Körperlänge 5.3-11.3 mm (Abb. 13), Kopfbreite 2.05-2.60 mm (Abb. 14). 

Diagnose d: Gelbe Zeichnungselemente gelegentlich reduziert; Quer- 
bänder auf dem Gaster meist durchgehend, Querband auf 1. Tergum nicht ein- 
geschnürt. 

Form des Gasters ähnlich L. dorsigera (Abb. 2b’). 

Körperlänge 7.0-9.2 mm (Abb. 13), Kopfbreite 2.10-2.40 mm (Abb. 14). 

Verbreitung: Paläarktisch, östlich bis Tadschikistan (Bouéek, 1974); in 
Mitteleuropa bisher nur aus der Schweiz (z. B. Schletterer, 1890; Madl, 1990), 
Tschechien, Slovakei (Bouéek, 1959) und Österreich (Madl, 1989) nachgewiesen. In 
der Schweiz ist das Vorkommen auf das Wallis beschränkt (Abb. 11), wo L. biguetina 
oft auch nach 1950 (57% der Daten, 69% der Fundpunkte auf der Verbreitungskarte) 
gesammelt wurde. Bemerkenswert ist das Fehlen von Daten aus dem unteren Teil des 
Wallis, d. h. unterhalb von Sion. Die Verbreitung stimmt recht gut mit derjenigen der 
potentiellen Wirte überein, wobei diese in der Schweiz weiter verbreitet sind (Nord- 
schweiz, GE, GR, TI) (Abb. 11). 

Höhenverbreitung: von 500 bis 1600 m ti. M. (Abb. 11). 

Gefährdung: gefährdet. Die beiden potentiellen Wirte (siehe unten) 
werden in der Roten Liste (Amiet, 1994) für die Südschweiz als gefährdet aufgeführt. 

Biologie: Aus Nestern von Osmia acuticornis Dufour & Perris (Le Goff, 
1996, 1997b sub Hoplitis) und O. tridentata Dufour & Perris (Le Goff, 1997a, 1997b 
sub Hoplitis) ın Frankreich gezogen. 

Imagines erscheinen von Juni bis September (Abb. 11) und werden normaler- 
weise an xerothermen Standorten wie Steppen- und Trockenrasen angetroffen. Ein- 
zelne Exemplare konnten auf Blüten von Doldengewächsen (Apiaceae) und Euphor- 
bia (Euphorbiaceae) beobachtet werden (vgl. Material; Amiet, Merz, pers. Beob.). 
Nach Bouéek (1974) vermehrt sich L. biguetina in Mitteleuropa parthenogenetisch, 
wobei aus unbefruchteten Eizellen fast durchwegs Weibchen entstehen (Thelytokie). 
Dies trifft wahrscheinlich auch auf die Population im Wallıs zu, da im reichen 
Untersuchungsmaterial von insgesamt 69 Exemplaren kein einziges Männchen vor- 
handen war. Allerdings scheint es sich hier um ein lokales Phänomen zu handeln, 
denn aus anderen, nahegelegenen Gebieten, z. B. dem Vinschgau im Südtirol, 
befanden sich unter insgesamt 9 Belegexemplaren 6 Männchen (vgl. Material). 


Leucospis bifasciata Klug, 1814 5a’, 5b', 12-14 


Untersuchtes Material: ASERBEIDSCHAN: Lenikoran, Azfilial Girkan Reserve 1 & (MH): 
NW Baku, Varafta Mts. 1 ¢ (MH). «Asia Minor: Namzur» 1 2 (ZSM). BULGARIEN: Slancev 
Briag 1 © (BZOL); SW Melnik 1 4 (BZOL). FRANKREICH: Basses-Alpes, Valensole 1 £ (FA), 
2 2 (NMBE). GRIECHENLAND: Kozani, Siatista, Brücke 1 £ (BZOL); Peloponnes, Zacharo 
1 2(ZSM); Preveza SB, 2 km S Kerasona 2 d (BZOL). ISRAEL: Antipatris 1 2 (ETHZ). 
JORDANIEN: N Shuna 1 4 (BZOL); Safi, 50 km of Karak 1 & (BZOL). ?LIBANON: «Sarepta» 
1 2(MHNG). SYRIEN: 20 km NE Latakia 1 2 (BZOL). TÜRKEI: Sultan Daglari, Yalvat 1 © 
(BZOL). TURKMENIEN: 15 km N Aschapat 1 2, 1 d (BZOL); Kopet-Dag Kizi-Arvat, 50 km S 
Chajagaia 1 2 (BZOL). USBEKISTAN: Yangikichlak, 10 km NW Ddjiak 1 d (BZOL). 


Diagnose 9 ähnlich der von L. dorsigera, aber: Scutellum am Hinterrand mit 
in der Mitte leicht eingeschniirtem Querband, dieses oft die Mitte des Scutellums 
iiberschreitend; Querband auf Gastraltergum 5 in der Mitte unterbrochen. 


382 HANNES BAUR & FELIX AMIET 


'.n=13____n=88 n=14_ n=186 _n=61___n=123_ n=79_n=18 
mig de (©) 


16— 


14= 


1.2 == 


1.0- 


©) 
Ar 
0.8— 
I I TT I I ra 
bifasciata brevicauda intermedia gigas Form 'A' 
biguetina dorsigera gigas Wallis gigas Form 'B' 


ABB. 12, Boxplot (nach Tukey) von Leucospis: Verhältnis Ovipositorscheide- zu Metafemur- 
länge. Die Grafik zeigt Median, Interquartilbreite, Variationsbreite (ohne Ausreisser), Aus- 
reisser (O), Extremwerte (%). 


Clypeus kaum vorgezogen und mehr oder weniger verrundet (Abb. 5a'); 
Flagellum stärker keulenförmig, Glieder 2-4 wenig kürzer als breit oder quadratisch; 
die beiden vorderen Querkiele des Pronotums kräftiger entwickelt als der hintere Kiel; 
Längsfurche des 1. Gastraltergums an der Basis deutlich verjüngt (Abb. 5b’); Ovi- 
positorscheide kürzer, 0.90-1.05 mal so lang wie der Metafemur (Abb. 12), reicht 
höchstens bis zur Mitte des 1. Tergums. 

Körperlänge 6.3-9.0 mm (Abb. 13), Kopfbreite 1.85-2.60 mm (Abb. 14). 

Diagnose d: Gelbe Zeichnungselemente gelegentlich reduziert; Quer- 
bänder auf dem Gaster normalerweise durchgehend. 

Form des Gasters ähnlich L. dorsigera (Abb. 2b’). 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 383 


Weibchen 
_n=13 __.n=88 __n=14_ _n=187 __n=61 n=123__ n=79 _ 


16= 
14= 
O 

| O 
12— 

O 

| o 
10— MES È TE 


PE 
8— — 

i | | | 

(Sra did 0 


6= na ce 


O Ben 
(©) 
bifasciata brevicauda intermedia gigas Form 'A' 
biguetina dorsigera gigas Wallis gigas Form 'B' 
Mannchen 
nn MENS, n26 1e NES6 n=10... n=42 
14— 
25, 
| o 
10— 
wi 
8— F E: I 
A — A 
| | 
6= pela 
bifasciata biguetina brevicauda dorsigera gigas intermedia 


ABB. 13, Boxplot von Leucospis: Körperlänge in mm. 


Körperlänge 4.7-8.3 mm (Abb. 13), Kopfbreite 1.40-2.30 mm (Abb. 14). 
Verbreitung: Paläarktisch, östlich bis Tadschikistan (Bouëek, 1974); in Mittel- 
europa wurde L. bifasciata bisher noch nicht nachgewiesen. Der Schweiz am nächsten 


384 HANNES BAUR & FELIX AMIET 


gelegene Vorkommen: Frankreich, Basses Alpes, Valensole (FA, NMBE); Italien, 
Piemont, Oulx und Veneto, Venedig (Boucek, 1974). 

Biologie: Aus Nest von Anthidium strigatum (Panzer) in Kroatien ge- 
zogen (Fahringer, 1922); Bestimmung des Parasitoiden überprüft von Bouëek (1974). 


Leucospis brevicauda Fabricius, 1804 2a-b, 12-14. 


Untersuchtes Material: ALGERIEN: El Afroun 1 d (MHNG). ITALIEN: Puglia, Mt. 
Gargano, Cagnavo 1 © (BZOL); Sardinien, Siniscola 1 £ (FNSM). MAROKKo: 4 2 (MHNG); 
70 km N Agadir, Tamri 1 © (BZOL); 100 km E Bouzakame, Icht | d (BZOL); Tanger 4 9, 
3 6 (MHNG); 10 km S Taroudant 1 4 (BZOL). TUNESIEN: Gabès, Skhira 2 2 (MH); Gafsa-El 
Guetar 1 2 (BZOL); Metouia 1 d (MH). 

Diagnose ® ähnlich der von L. dorsigera, aber: Flagellum rötlich; 
Zeichnung neben schwarz und gelb auch rotbraun; gelbe Zeichnung häufig stärker 
ausgedehnt als bei L. dorsigera, besonders an Pronotum, Mesoscutum, Scutellum und 
Gaster; Metafemur ausser an den Zähnen und an der Basis gelb (Abb. 2a); Metatibia 
dorsal rotbraun bis schwarz, höchstens in der Mitte mit feinem gelblichem Längs- 
streifen; Tarsen meist rotbraun; Gastralterga 1, 4, 5 und 7 mit variablen Querbändern, 
auf den Terga | und 4 manchmal, auf Terga 5 und 7 immer in der Mitte unterbrochen; 
5. Tergum meist zusätzlich mit kleinen Flecken; auf dem 7. Tergum helle Zeichnung 
rotbraun, daher oft undeutlich. 

Clypeus kaum vorgezogen und mehr oder weniger verrundet (ähnlich L. 
bifasciata, Abb. Sa'); die beiden vorderen Querkiele des Pronotums kräftiger ent- 
wickelt als der hintere Kiel; basaler Zahn des Metafemurs ungefähr so lang wie 
folgende Zähne (Abb. 2a); Gastralterga 1-4 ohne Längsfurche (ähnlich L. biguetina, 
Abb. 4b), diejenige des 5. Tergums tief, so dass Ovipositorscheide in Ruhelage 
teilweise verdeckt ist (ähnlich L. biguetina, Abb. 4c); Ovipositorscheide sehr kurz, 
0.66-0.74 mal so lang wie der Metafemur (Abb. 12), reicht nur bis zur Mitte oder zum 
basalen Drittel des 5. Tergums. 

Körperlänge 5.8-9.2 mm (Abb. 13), Kopfbreite 1.90-2.45 mm (Abb. 14). 

Diagnose d: Gelbe Zeichnungselemente gelegentlich reduziert; 
Querbänder auf dem Gaster durchgehend. 

Gaster 1m basalen Drittel stark verschmälert (Abb. 2b). 

Körperlänge 6.8-8.2 mm (Abb. 13), Kopfbreite 1.45-2.15 mm (Abb. 14). 

Verbreitung: Circum-mediterran (Bou,,ek, 1974); in Mitteleuropa 
wurde L. brevicauda bisher noch nicht nachgewiesen. Der Schweiz am nächsten 
gelegene Vorkommen: Frankreich, Hérault, Palavas (Berland, 1934b) und Bouches- 
du-Rhöne, Port du Bouc (Bouëek, 1974): Italien, Puglia, Mt. Gargano (BZOL). 

Biologie: Wirt unbekannt. 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 385 


Weibchen 

__n=13 __n=88 _n=14 «  n=187 n=61 n=123 n=79 n=18 
4.0— 
3.5= | 


oi le —- ape 


a 
A 
TH 


IS Q 


bifasciata brevicauda intermedia gigas Form 'A' 
biguetina dorsigera gigas Wallis gigas Form 'B' 
Männchen 
Zn INT m eee Ai Ve EURE a 
3.0— calli 
O — 
2.5= | 
©) 
2.0— Pre wa 
A 
1.5— 
© 
bifasciata Core Poe dorsigeran gigas intermedia 


ABB. 14, Boxplot von Leucospis: Kopfbreite in mm. 


386 HANNES BAUR & FELIX AMIET 


DANKSAGUNG 


An erster Stelle gebührt unser Dank Elsa Obrecht (NMBE) für die sorgfältige 
Durchsicht des Manuskriptes sowie die Ausleihe des Materials am NMSO. Rene 
Hoess, Bern, Schweiz, las eine frühere Version des Textes, und wir verdanken ihm 
viele nützliche und kritische Anregungen. Wir danken ebenfalls einem anonymen 
Rezensenten. Charles Huber (NMBE) hatte entscheidenden Anteil bei der Erstellung 
der Verbreitungskarten, für die Geduld gegenüber den vielfältigen Wünschen des 
einen Autors (H.B.) sei herzlich gedankt. Bertrand Baur, Rumendingen, Schweiz, 
begleitete H.B. auf einer Exkursion ins Wallis und machte Habitataufnahmen. Ingmar 
Wall (IW) überliess uns bereitwillig seine Notizen zu einem Bestimmungsschlüssel 
der europäischen Leucospis-Arten. Frank Burger (FB), Jakob Forster, Winterthur, 
Schweiz, Gerd Reder (GR), Albert Sermet (ASE), Erwin Steinmann (ES), Hansueli 
Tinner-Guler (HT) und Jürg Zettel, Zoologisches Institut, Bern, Schweiz, teilten uns 
ausführlich Freilandbeobachtungen zum Verhalten einiger Arten mit und stellten 
zusätzlich eigenes Sammlungsmaterial bzw. hervorragende Dias zur Verfügung. 
Ihnen allen sei herzlich gedankt. Für die Ausleihe von Material oder Angaben zu 
Exemplaren in Sammlungen danken wir schliesslich den folgenden Personen und 
Institutionen: Werner Arens (WA), Stefan M. Blank (DEI), Hermann Blöchlinger, 
Naturmuseum Thurgau, Frauenfeld, Schweiz, Zdenék Boucek, The Natural History 
Museum, London, UK, Daniel Burckhardt (NMBA), Toni Bürgin, Naturmuseum, St. 
Gallen, Schweiz, Heinrich Bürgis (HB), Gilles Carron (GC), Erich Diller (ZSM), 
Emanuel Gerber, Musée d’histoire naturelle, Fribourg, Schweiz, Fritz Gusenleitner 
(BZOL), Martin Hauser (MH), Ulrike Hausl-Hofstätter, Landesmuseum Joanneum, 
Graz, Österreich, Peter Huemer, Tiroler Landesmuseum Ferdinandeum, Innsbruck, 
Österreich, Frank Koch (MNHU), Jens-Peter Kopelke (FNSF), Manfred Kraus, 
Nürnberg, Deutschland, Walter Linsenmaier (WL), Ivan Löbl und Bernhard Merz 
(MHNG), Paul Mildner, Landesmuseum für Kärnten, Klagenfurt, Österreich, Andreas 
Müller (ETHZ), Till Osten (SMNS), Enrico Ratti (MSNV), Helmut Riemann, 
Ubersee-Museum, Bremen, Deutschland, Ali Salvioni (AS), Michel Sartori (MZL), 
Christian Schmid-Egger (CS), Stefan Schödl (NMW), Ingmar Wall. 


LITERATUR 


AMIET, F. 1994. Rote Liste der gefährdeten Bienen der Schweiz (pp. 38-44). In: Duelli, P. (ed.). 
Rote Liste der gefahrdeten Tierarten der Schweiz. Bundesamt fiir Umwelt, Wald und 
Landschaft, Bern, 96 pp. 

BERLAND, L. 1934a. Un cas probable de parthénogenèse géographique chez Leucospis gigas 
(Hyménoptère). Bulletin de la Société Zoologique de France 59: 172-175. 

BERLAND, L. 1934b. Notes sur les hyménoptères. XVI. Les Leucospis de France (Chalcididae). 
Revue française d'Entomologie 1: 65-69. 

BOUCEK, Z. 1959. A revised key to the West-Palearctic species of Leucospis (Hym. Chalc.), 
with some new synonymy. Acta Entomologica Musei Nationalis Pragae 33: 435-444. 

BOUCEK, Z. 1964. Leucospidae (pp. 10-11). n: Peck, O., Bouéek, Z. & Hoffer, A. (eds): Keys 
to the Chalcidoidea of Czechoslovakia (Insecta: Hymenoptera). Memoirs of the 
Entomological Society of Canada 34: 3-120. 

Boucek, Z. 1970. Contribution to the knowledge of Italian Chalcidoidea, based mainly a on 
study at the Institute of Entomology in Turin, with descriptions of some new European 
species (Hymenoptera). Memorie della Societa Entomologica Italiana 49: 35-102. 


LEUCOSPIDAE (HYMENOPTERA) DER SCHWEIZ 387 


Boucek, Z. 1974. A revision of the Leucospidae (Hymenoptera: Chalcidoidea) of the world. 
Bulletin of the British Museum (Natural History) Entomology, Supplement 23: 1-241. 

BOUCEK, Z. 1977. A faunistic review of the Yugoslavian Chalcidoidea (parasitic Hymenoptera). 
Acta entomologica Jugoslavica 13, Supplementum: 4-145. 

BÜRGIS, H. 1995. Leucospis gigas (Chalcidoidea: Leucospidae) als Parasit der Mörtelbiene 
Megachile sicula (Apoidea: Megachilidae). Bembix 5: 27-32. 

BYTINSKI-SALZ, H. 1963. Geographical variation and sex-ratio in Leucospis gigas (Hyme- 
noptera, Chalcidoidea). Acta Entomologica Musei Nationalis Pragae 35: 527-530. 

CALECA, V., LO VERDE, G. & Massa, B. 1995. Leucospis miniata Klug a Lampedusa 
(Hymenoptera Chalcidoidea Leucospidae). /n: Arthropoda di Lampedusa, Linosa e 
Pantelleria (Canale di Sicilia, Mar Mediterraneo). Naturalista siciliano 19 
(Supplementum): 773.775. 

CHEVRIER, F. 1870. Description de quelques Hyménopteres du Bassin du Léman. Mitteilungen 
der Schweizerischen Entomologischen Gesellschaft 3: 265-276. 

CLAUSEN, C. P. 1940. Entomophagous insects. New York, X + 688 pp. 

FABRE, J.-H. 1886. Souvenirs entomologiques (troisième série). Paris, 329 pp. 

FAHRINGER, J. 1922. Beiträge zur Kenntnis der Lebensweise einiger Chalcididen. Zeitschrift für 
wissenschaftliche Insektenbiologie 17: 7-13, 41-47. 

FUESSLY, J. C. 1783. Leucospis dorsigera Fabr. Ein Beytrag zur nähern Kenntnis dieses Insects. 
Archiv für Insectengeschichte 3: 1-2, Tafel 18. 

GIBSON, G. A. P. 1997. Morphology and terminology (pp. 16-44). In: Gibson, G. A. P., Huber, 
J. T. & Woolley, J. B. (eds). Annotated keys to the genera of Nearctic Chalcidoidea 
(Hymenoptera). NRC Research Press, Ottawa, Ontario, Canada, 794 pp. 

GIRAUD, J. 1858. Note sur un hyménoptère nouveau du genre Ampulex trouvé aux environs de 
Vienne. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Osterreich 8: 441- 
448. 

GIRAUD, J. E. & LABOULBENE, A. 1877. Liste des éclosions d'insectes. Annales de la Société 
entomologique de France 7: 397-436. 

GOULET, H. & Mason, W. R. M. 1993. Use of keys (pp. 60-64). Jn: Goulet, H. & Huber, J. T. 
(eds). Hymenoptera of the world: An identification guide to families. Canada Commu- 
nications Group, Ottawa, Canada, VII + 668 pp. 

GRANDI, G. 1961. Studi di un entomologo sugli imenotteri superiori. Bollettino dell’Istituto di 
Entomologia dell'Universita di Bologna 25: 1-659. 

HAGENBACH, J. J. 1822. Symbola Faunae Insectorum Helvetiae, exhibentia vel species novas 
vel nondum depictas. Fasciculus Primus. Basel, VI + 7-48 pp. + 15 Tafeln. 

HERTING, B. 1973. Coleoptera to Strepsiptera. A cataloque of parasites and predators of terres- 
trial arthropods. Section A. Host or prey/enemy. Commonwealth Agricultural Bureaux, 
218 pp. 

HERTING, B. 1977. Hymenoptera. Part 4. A cataloque of parasites and predators of terrestrial 
arthropods. Section A. Host or prey/enemy. Commonwealth Agricultural Bureaux, II, 
206 pp. 

JacoBs, W. & RENNER, M. 1998. Biologie und Ökologie der Insekten: ein Taschenlexikon. 3. 
Auflage, tiberarbeitet von K. Honomichl. Fischer, Stuttgart, 678 pp. 

JURINE, L. 1807. Nouvelle méthode de classer les hyménoptères et les diptères. Hyménoptères. 
Tome premier. Genève, 320 pp. + 14 Tafeln. 

KLUG, F. 1814. Die europäischen Arten der Insekten-Gattung Leucospis. Magazin der Gesell- 
schaft naturforschender Freunde zu Berlin 6: 65-71. 

LABRAM, J. D. & IMHOFF, L. 1836. Insekten der Schweiz, die vorzüglichen Gattungen je durch 
eine Art bildlich dargestellt. Erstes Bändchen. Basel, unpag. 

LAUBER, K. & WAGNER, G. (1996): Flora Helvetica. Haupt, Bern, 1613 pp. 


388 HANNES BAUR & FELIX AMIET 


LE Gorr, G. 1996. Découverte d’un hôte de Leucospis biguetina Jurine (Hymenoptera Chalci- 
doidea Leucospidae). L’Entomologiste 52: 62. 


LE Gorr, G. 1997a. Addendum à la note relative au site d'Opoul (Pyrénées Orientales) sur les 
nidifications d'apoïdes et la présence de Leucospis biguetina Jurine (Hymenoptera). 
L'Entomologiste 53: 28. 

Le Gorr, G. 1997b. Note sur la nidification dans la férule d’apoides Anthophoridae et Mega- 
chilidae du Roussillon (Hymenoptera - Apoidea). L’Entomologiste 53: 259-269. 

LE GOFF, G. 1999. Leucospis dorsigera Fabricius, parasite d'Osmia (Osmia) tricornis Latreille 
(Hymenoptera). L’Entomologiste 55: 89-90. 

Map , M. 1989. Zur Kenntinis der paläarktischen Leucospis-Arten unter besonderer Berüch- 
sichtigung der Fauna Osterreichs (Hymenoptera, Chalcidoidea, Leucospidae). Entomo- 
fauna 10: 197-201. 

MADL, M. 1990. 2. Beitrag zur Kenntnis der paläarktischen Leucospis-Arten unter besonderer 
Berüchsichtigung der Fauna Österreichs (Hymenoptera, Chalcidoidea, Leucospidae). 
Linzer biologische Beiträge 22: 81-87. 

MULLER, A., KREBS, A. & AMIET, F. 1997. Bienen: Mitteleuropäische Gattungen, Lebensweise, 
Beobachtung. Naturbuch, Augsburg, 384 pp. 

Noyes, J. S. 1998. Catalogue of the Chalcidoidea of the world. Biodiversity catalogue database 
and image library CD-ROM series. ETI Biodiversity Center, Amsterdam, Netherlands. 

PAGLIANO, G. 1998. Leucospididae d’Italia e indagine sulla loro presenza in Piemonte 
(Hymenoptera, Chalcidoidea). Rivista Piemontese di Storia naturale 19: 247-258. 

RONDANI, C. 1873. Degli insetti nocivi e dei loro parassiti. Enumerazione con note. Bollettino 
de la Società Entomologica Italiana 5: 209-232. 

SAUNDERS, S. S. 1875. Exhibitions, & c. Proceedings of the Entomological Society London 
1875: XVII [Im Anhang zu Transactions of the Entomological Society London 1875, 
Teil 5]. 

SCHLETTERER, A. 1890. Die Gruppe der Hymenopteren-Gattungen Leucospis Fab., Polisto- 
morpha Westw. und Marres Walk. Monographie. Berliner Entomologische Zeitschreift 
35: 141-302 + 2 Tafeln. 

ScHMID-EGGER, C. 1995. Die Verbreitung von Leucospis dorsigera in Deutschland (Chalci- 
doidea). Bembix 4: 6. 

SCHMIDT, K. 1969. Beiträge zur Kenntnis der Hymenopterenfauna des Mittelrheingebietes, 
insbesondere des Mainzer Sandes. Mainzer naturwissenschaftliches Archiv 8: 292-302. 

SCHMIEDEKNECHT, O. 1930. Die Hymenopteren Nord und Mitteleuropas. Fischer, Jena, 1062 pp. 

SCHWARZ, M., GUSENLEITNER, F., WESTRICH, P. & DATHE, H. H. 1996. Katalog der Bienen 
Osterreichs, Deutschlands und der Schweiz (Hymenoptera, Apidae). Entomofauna, 
Supplement 8: 1-398. 

SULZER, J. H. 1776. Abgekürzte Geschichte der Insecten. Nach dem Linnaeischen System. Teil 
1 + 2. Winterthur, 274 pp. + 32 Tafeln. 

WESTRICH, P. 1989. Die Wildbienen Baden-Wiirttembergs. Allgemeiner Teil. Ulmer, Stuttgart, 
431 pp. 

WESTRICH, P. & DATHE, H. H. 1997. Die Bienenarten Deutschlands (Hymenoptera, Apidae). 
Ein aktualisiertes Verzeichnis mit kritischen Anmerkungen. Mitteilungen Entomolo- 
gischer Verein Stuttgart 32: 3-56. 

WOLF, H. 1953. Beiträge zur Hymenopterenfauna des oberen Lahn-Dill-Sieg-Gebiets III. 
Mitteilungen des naturwissenschaftlichen Musums Aschaffenburg 41: 83-85. 


REVUE SUISSE DE ZOOLOGIE 107 (2): 389-418; juin 2000 


The Bufo tuberosus species group with the description 
of a new species from the rainforest of Cöte-d'Ivoire 


Mills TANDY! & Jean-Luc PERRET? 

! Colorado State University 
201 Highland Drive 
Sierra Vista, Arızona 85635 USA. 

? Département d’herpétologie et d’ichtyologie 
Muséum d’histoire naturelle 
Route de Malagnou 1, case postale 6434 
CH-1211 Geneve 6, Suisse. 


The Bufo tuberosus species group with the description of a new species 
from the rainforest of Cöte-d’Ivoire. - A very small species of toad 
similar to Bufo tuberosus was discovered in a collection made in West 
Africa by M. Lamotte in the late 1960s. In the 1980s a second specimen of 
this species from Cöte-d’Ivoire was donated to the Muséum d’histoire 
naturelle de Geneve. This paper describes the new species, B. amieti, 
presents additional data on B. tuberosus, compares these species to others 
and hypothesizes an origin of B. amieti and B. tuberosus by geographical 
isolation of a common ancestor in eastern and western blocks of equatorial 
rainforest separated by the V-Baoulé during past interpluvial periods. 


Key-words: Bufo - Bufo tuberosus group - speciation - Céte-d’ Ivoire - rain 
forest - West Africa - Anura. 


INTRODUCTION 


This paper summarizes the biology of the central African rainforest toad Bufo 
tuberosus Günther and describes a new species of the same group from the far 
western section of African equatorial lowland rainforest in Côte-d'Ivoire. 

Bufo tuberosus, a very distinctive species from the rainforests of central 
Africa, was described in the mid nineteenth century. It is so morphologically different 
from other African Bufo with its usually globose parotoid glands and extreme 
development of spinose warts that it was placed in its own species group (Tandy, 
1972; Tandy & Keith, 1972). Although Bufo tuberosus was described very early in 
the literature on African amphibians, and collections have been made by various 
investigators over the years, it is not a common amphibian in museum holdings. 
Little is known about its behavior and nothing about its genetics. This paper analyzes 


Manuscript accepted 08.01.2000 


390 MILLS TANDY & JEAN-LUC PERRET 


morphological variation based on museum material, reports observations of its calling 
behavior and recordings of ıts mating call made in Camerounian rainforests by Amiet 
(1973, 1975, 1976, 1989) and by Tandy & Tandy (1976) and considers ecological 
features of known localities. 

In the 1970’s, Tandy encountered a small toad in a collection from Graba- 
zouo, Côte-d'Ivoire made by M. Lamotte and R. Vuatroux. This is a gravid female 
that resembles Bufo tuberosus but which is half the size of adult females of that 
species. In 1986, J.-L. Perret received a second female of this species, collected by R. 
Neumeyer, from Tai, Cöte-d’Ivoire. This paper describes these specimens as a new 
species and compares them to Bufo tuberosus and other species from central and 
western Africa. The new species is named in honor of our colleague who has contri- 
buted so much to central African herpetology, J.-L. Amiet. 

Terminology and methods of data collection and analysis are given in Tandy 
(1972), Tandy & Keith (1972), Tandy & Tandy (1976), Largen et al. (1978), Tandy et 
al. (1982), Tandy er al. (1985) and Tandy & Feener (1985). 

Data are presented in the following format: locality, geographic coordinates; 
sex and collection numbers of specimens; collection numbers of tape recordings; alti- 
tude; date of collection; collector. 

The following abbreviations are used for collection numbers: 


AMNH American Museum of Natural History, New-York; 


BM British Museum (Natural History), London; 
CAS California Academy of Sciences, San Francisco; 
CNHM Chicago Natural History Museum, Chicago; 
FM Field Museum of Natural History, Chicago; 


MBG Mission Biologique au Gabon; 

MCZ Museum of Comparative Zoology, 

Harvard University, Cambridge, Massachusetts; 

MHNB Muséum d’histoire naturelle, Bale; 

MHNG Muséum d’histoire naturelle, Geneve; 

MHNCF Muséum d’histoire naturelle, La Chaux-de-Fonds; 

MNHP Museum national d’histoire naturelle, Paris; 

MT Collection of M. Tandy; 

MT() Reference number of M. Tandy assigned to specimen of collection 
indicated within parentheses; 

MT.TC.Ca Collection of tape recordings of M. Tandy; tape cut from Cameroun; 

RGMC Registre général du Musée du Congo, Tervuren; 


SMF Natur-Museum und Forschungsintitut Senckenberg, Frankfurt; 

UMMZ Museum of Vertebrate Zoology, University of Michigan, Ann Arbor, 
Michigan; 

ZFMK = Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn; 

ZMB Zoologisches Museum der Humboldt Universitat, Berlin. 


Bufo amieti sp. n. 


Holotype: 1 adult female (MHNG 2594.17) from lowland rainforest near Tai, Tai Na- 
tional Park, Côte-d'Ivoire (5° 52° N 7°28’W, altitude 123m) collected in 1986 by R. Neumeyer. 


THE BUFO TUBEROSUS SPECIES GROUP 39] 


Fic. 1. Bufo amieti sp. n. Holotype female (above): habitus, natural size; (below): detail of the 
enormous paratoid gland, tympanum and eye. 


Paratype: 1 adult female (MNHP 1997 4934 = MT(P) 6141) from lowland rainforest 
on the Sassandra River near Grabazouo, Soubré, Cöte-d’Ivoire (5° 50’N 6° 35°W, altitude 
~150m) collected 10.XII.1967 by M. Lamotte and R. Vuatroux. 

Diagnosis 

A very small species of Bufo resembling Bufo tuberosus Giinther, 1858, but 
only half the size of that species. Body measurements and their ratios indicate at least 
five diagnostic differences with all species to which B. amieti has been compared 


392 MILLS TANDY & JEAN-LUC PERRET 


Fic. 2. Bufo amieti sp. n. Holotype female: dorsal and ventral color pattern. Magnification 2X. 


THE BUFO TUBEROSUS SPECIES GROUP 393 


Fic. 3. Bufo amieti Sp. n. Paratype female: dorsal and ventral color pattern. Magnification 2X. 


394 MILLS TANDY & JEAN-LUC PERRET 


Fic. 4. Bufo amieti sp. n. female (paratype: MHNP 1997 4934); snout-urostyle length 35.9 mm. 
Grabazouo, Soubré. Côte d'Ivoire; Bufo tuberosus female (BM 1909.7.9.14); SU 65.8 mm. 
Efangono, Cameroun; B. tuberosus male (BM 1934.12.1.14); SU 45.0 mm. Lomié District, 
Cameroun. 


(Tables | - 3). Comparisons have been made with all species of Bufo known from the 
geographic area concerned, west and central Africa, and with all members of the Bufo 
regularis Complex because of the acoustical similarities between the mating call of 
Bufo tuberosus and species of that complex. The ratio of tympanum diameter/parotoid 
width separates B. amieti from almost all congeners. Tympanum diameter, snout- 
urostyle length or parotoid length/parotoid width distinguish it from the others. 
Tympanum distinct, its horizontal diameter approximately 0.12 the width of the head 
and 0.18 the width of the parotoid; snout-urostyle length about 37mm; parotoid length 
1.7 times the parotoid width; no tarsal fold; male unknown; first finger longer than 
second; toes not extensively webbed (33/4 phalanges of fourth toe free of webbing on 
both inner and outer margins); lacking series of warts on posterior surface of forearm; 
large warts with multiple cornified spinules extensively developed. Ecological data 
for two localities: lowland rain forest, altitude, 


X = 136.5m, range (123-150); mean annual rainfall, 
x = 173.70cm (161.1-186.3); mean annual temperature, x = 23.95°C (23.9-24)(Table 4). 


DESCRIPTION 


Holotype: Snout-urostyle length (SU) 37.7mm; form moderately stout; head 
triangular, broader than long, head width (HW) 15.5mm, head length (HL) 10.2mm; 
snout obtusely rounded; nostrils closer to tip of snout than to eye; canthus rostralis 


395 


THE BUFO TUBEROSUS SPECIES GROUP 


te] ve 0 SL'9 GOP MSG EI EEE SIE LG [RC 
OT | LIO DIE OTO OFT ISLAS SI IST 661 
STIOO 98000 SSOEO TETOO LOOTO ETSTO S600 0 SErO'O 0LEOO OILO 
LSTI. VOTO OCTO PEED COBT 9097 0810 9IÿT CSLT OST 
LET 8c 0 GOT = 8Si0r 88567 SS9 OT ON ETAT OS E 0€ 
PIT eco 89'€ 8CO  6rl rl EIEIT IEG ECO VVC 
11700 OF00'0 SIZ90 6ÿc00 9EITO S66€ 0 8E00'O ZLEPO'O TOr0'0 E6S O0 
OSTT CSTO 9996 9€€0 LIST 980E 98T0 L6PI COST O9LT 


tel ££'0 LS LORS ET eee 0 VETO L'6€ 
sol 810 OLT GO) tS || cel TiO 9a Sexe r’87 
ISTO loo L00°0 9700 
COTE 1670 stIO LLOT 


trl bro LOT DO CHIC ee SO Velie NTTIE 9'£9 
IUT LT'O SET CO Hr LOT ITO OCT OST 


cs1I00 I0100 88/00 92/00 0 097000 S6E00 CE00 0 81r0°0 F6co'0 8760 
VITI SCO 6ETE LLTO L661 9€91 6€TO SOLI 69LT 860$ 
dal OT'0 81€ GIORIO ETA TAO COTE 206i SIT 
or I LIO 850 610 191 OST ONCE CET 9'CC 
CTITT €810 1887 9610 LSOT 9ELT GITO SEST 1997 S097 
dal OT0 857 005 Sele 09M COS CONTE erase SIT 
OT I LIO ste [TO OLI L481 CIO SST 067 IST 
[sey 


HO/NS = Md/L = Md/MH Id/L ‘Id/MH Md/Id 


cr OL vv 
OT Ut VC 
90T0 =LSEO 


UA? 18 97 
I OV UE 
r610 HEEO 


(GE) 001 c9 
SE vo se 
£TTO SSEO 


S'6 PSI cs 
vs 6 St 
0970 


£OT 0 
rec ILS 81€ 


8IT0 


MAE Vo) == 1s 


ILTO 
SOTA IE sort 


9£T'O 
LE CT 239 


9T 
Ir1°0 
SCT 


OV 
VC 
1600 
6l’e 


CEI 
98 
Loc 0 
re Ol 
gel 
PTT 
ECCO 
SEI 


107 
Vel 
809°0 
€9 SI 


997 
€ 61 
ece 0 
PS TC 


8 9€ 
BET 
9¢0'1 
6C'8T 
8 LE 
VIE 
065°0 
8STE 


l'8r 
8 CE 
ler] 
OT'6€ 


UNULIXEJA] 
WINUILUIN 


Jong prepurys 
LI SEUL - uroW 


wnumxeW 
umwmu] 


10119 prepurys 
I] sopewaJ - uraW 


uUogeH ‘UNOIAUP) 
sadij19043 ofng 


wnwıxeW 
umwmu 

IOLA prepuris 

II sopew - BON 


[elio eno vauiny ‘unosoweD 


VIL 
evs 
13220) 
IIE9 


umwmxeWN 
umwmu 

Jong pIepuels 

je SU] - ueaW 


o8u0) np anbne.wowog anbipgnday “TeLIoensg vous ‘“uno1sumry 


9 96 Ir 
6€ Gil ve 
Sor St8 SL’ 
09 96 Ir 
6€ el ve 


wWpIMm Yu] JOJOUPIP 19JIUILIP 
MH/L TH/MH MH/NS -PIAsoan proyeieg plowieg Ag wnuedwA] pray 


Sup] 


SSI 
rc 
SG ET 
SSI 
ro 


pin 
peoH 


ESL 
6 SE 
8 9€ 
LUE 
6SE 


2[41s01n 
-jnous 


snsosaqny ofng 


wnuxeW 
wnunumN 
T SO[euI9] - uvoW 
BL 
ONOZEgGRIO 


SIIOALP 9199 
yanup ofng 


u 


"ww ur 918 SJUDUDINSKILU [[Y ‘YTU] [eoy-a]A}soin = HN ‘YIPIM projored 
= Md ‘sua piojosed = "Id ‘249 ay) Jo YPPWEIP [EJuOZLIOy = 4 ‘WunuedwA} OY} JO 1SY9WweIP [RIUOZLIOY = L ‘Yus] PEAU = TH ‘UIPIM peau = 
MH ‘8ua] a[Ajsoin-jnous = NS ‘POLY [HU pur Is9M WO Sa19ods 19yJo pur 2722242 ofng 107 SONCI Nay) pur SJUSWSANSEIU IOUT] “| ATAVL 


MILLS TANDY & JEAN-LUC PERRET 


396 


oc I 
LOI 
s110'0 
ss 


OE I 
811 
£9100 
9ET I 


HO/NS 


ei 
L9'0 
10E0'0 
9960 


vel 
09°0 
chS0°0 
reo 0 


cL 0 
sp 0 
Ir10'0 
EOIO 
cs 0 
GEO 
ELVO 


6€ 0 
dat) 
TICO 
bro 
ICO 
eIEO 


Md/L 


po'8 
ssp 
Ore9 


AR 
61+ 
LLV1 0 
OSTS 


LINE 
WS 
SI6T 0 
CLES 


sp} 
[CE 
9190 0 
pele 
8S 
68°C 
IITE 


9ST 
spe 
LSG'E 
00? 
LA 
LELE 


Md/MH 


Oe 0 
OTO 
Irc'O 


ro 
9070 
07100 
She 0 


170 
9T 0 
911000 
LAGO) 


IEO 
610 
6800 0 
9LTO 
red 
810 
IICO 


ICO 
LIO 
SETO 
ce O 
STO 
IETO 


Id/L 


Fl Vip — LO 
hil We CO 
9591 OSL'E 9FL-0 
One “GES 2 460 
ari dle ORO 
F050'0 86600 LEOO'O 
SS8T G6LT 8810 
SoC le Koll) 
6 600 no 
LZ90'0 ISPI’O FE00'0 
9981 0687 FLI'0 
VIT 987 610 
kei Ser. SIRO 
FLr0'0 FE90'0 LZ00'0 
GTLI OCTT 6S1°0 
US. SPE SO 
GEL CE IO 
LOT IPTC Pro 
SLT 097 810 
HI Of€% O10 
E191 St 9FI‘0 
307 9% 910 
671 TE O10 
EST PSC Trio 
IV/MH Md/Id MH/L 


VEG 09% 
Geil) BUG 
COLTI 9LV'C 
86 I CLIC 
OG - OS 
0090 0 LLI0'0 
CUE, GS 
oT 195% 
CSI ICT 
csr0 0 £9CO'0 
GGL GEV 
LS I COC 
OG 99 
£8100 SSIO0 
GLE SOLE 
S9 I SLT 
CE WEG 
08F 1 tcl 
O8 T 90€ 
CS I 89 T 
STI 6787 
60c LOE 
VEIT 
crs I 9087 


TH/MH MH/NS -21AS0IN plored pion 


0'£9 
csr 
OSO I 
CS re 


Lee 
197 
CLEO 
tC GC 
121723 
9'O€ 
6 CE 


OSs 
Ot 
L8'tr 
19S 
PSE 
0667 


peau 


9Y 
OE 
£800 
COE 
vs 
sr 


O'S 


PM 


O00C 86 Ov 
ISIE Cy) 1883 
EN EE Olly 
VECI 405) Ot 
SL 8% OE 
ECAOGTROTLON 
LENO ES AOS te 
ysl #8 vs 
GLE GS) ve 
L8YO L810 9ELO 
BF EC 

LOI OS IT 
OS Iv LI 
£9T'O 1800 800 
r0'8 LLY 81°C 
O'CI O'S ST 

€ 0I cs IC 

€ II vs VC 


Orel 59 SE 
Cll 1S OTC 
(AT LOGS Ss 10 SEE 
CSI OL Ot 
SCI es 61 
Bose TEE 


SU] JOJAUUPIP 19JAUUPIP 
d 244 wnuedwX] 


SCT VVE L'E8 WNWIXENN 
CEI OSC 619 LUN LULU 
JOIN GAS CONTE 8  sopetluogf = UCSIN 
CLOU] ‘Q9UNO ‘LUPUO 'D.OALP 909 
SISUIUNIQUIDI Ofng 
8 SI SIC TSS WNWIXEN 
68 vgl OCP LUN LULA] 
IESO SO ISO JOLIE] P-IEPULIS 
SIIT STASI PI SOJELL = ULI 
[ELIO] NOT] BOUND "Tun OLY Ay UNO) 
PSI 1 DE Il LUNLUTXR A 
SCI OPC E 8S LUNLUTUTYAY 
(YES 9¢S:0 heal plepuris 
18 ST 80.7 6899 el SOUL] - UBOW 
OSUOD np anbıpgnday ‘eosin ‘UNOIOUP) 
SISUIUNAQULDI OfNG 
SII SI 444 LINKEN 
9'8 SII O'TE WINLUTUTN 
OLIO 6ET0 £690 FT piepuns 
766 GOIET QLYLE (U6 SO[UUI - ULOIN] 
SII CLI 89 LUNUIIXE]A 
66 gs] 6cr LLINLLILUTN 
II pol Yrr € SAW) - UPON 
AOA], P 9109 ‘PÉLIOUOIN 
apjalupp ofng 
CCI S'OT Soc WNWIXEN 
801 08T  Oips LUNUUN 
CO ‘8061 SEHE 9 sojeu - UVOIN 
OSI 09T 0'069 WINKEN 
Sol lol E CS LUN LULU 
vycl €8¢C LEED L  So[PUO] - uray 
UNOIOUUPE) 
18421]]1A Ofng 
yısus] ppia u 
peoH proH 


37 


THE BUFO TUBEROSUS SPECIES GROUP 


Soul sol T9 L CGONTOICMETENTTIONNE CAT Sec 697 8% AGIT CHE SC CET SIT OS umuxeN 
OT] cS0 Ich LIO Lv LEVG CO AZ 16€ IT L'6 es EIT vII MSI € Ly WUNUITUTIN 


CLI STO)” (88S ISIGION (SYV9ANE See. OEKO LS LIE I SE SOI Vous AO COS EST gETI 6761 S8SSIS 0'8 Sa[UU - UEOWN 


PE €60 €89 SCOR aL al eer +10 0671 19°C 19 LA 9IT 68 6€ L'OT L'8T OIL UNUUTXEIN 

STI 00 VIE POST [ST ÉNOMECON MERS CT Tes CV CLI iY) l'E VPI LYC 8 19 WUNUUTUTIN 

ICT NOTA “98S (8810 TOITS TL IE 6610 SL STE MEDI ARTI CSST 0G" (OS SGSW Enke 0189 Oy SITES] - USW 
030L 

SISU2050] ofng 

STI LST OUI CENT E80 20 0; EIKON IO SEVEN, SOI UL SC Or IIC L'SS UUNWITXE INT 

860 tr 0 607 610 661 981 80/0} MET SO CVE VI SO vv SI € 6 € SI L'6€ WNUNUTN 

ccloOO GESOO EOFEO [8000 CI900 [0610 9£000 6PE0°0 99€0 0 9ELO LPO €600 srlO cLOO ILTO ITEO 0580 Joly piepue]s 

SIETEESIAOEFELEITESIENTZLIETETBESITTTT ET BEE IT EEG CH LCV. {Olen a LOS CONSE Se Gln LLIT v88l cr 6r EC Ssojeu - USM 
uno.) 


suo4fnp] ofng 


cc I 680 LA CAOS 09 EROE CI 06 8 €9 vs 9'CI SL Gis L'LT GST (Gale WUNUXEIN 
CI 050 PST VASO IONI WORST SEE ers 87 CE 6 89 ST PSI 8 CC c 99 UAUUTUTIN 
IS CIAO! 6969955 S9C0 VOR ZOO VSO! EST BIETET NOOBS EHE NOLO EEE 86ST £9ÿcC 80 69 y SOUS - UE 


[ELIOYENIT voUINy ‘TUNTA OLY a UNOIUWIEI 
suo.1f110] ofng 


sl] sc I 8CL PIO Tee RN O HR OIC OOS 8° GEI 89 O's OLI CIE CMS UMTUTXEIN 

[FI cL'0 COLE 600. 191 OCT 80 VE 68T Vice SS 159 LS TE 601 CSI VSP UAUTUTIN 

LEVIN 25660 “970: 7EIEO VLvol 70785 200407 BEI COSCA LEViOne COSTA (Stall G9, “Loire © 89M TNT 05 9 SOJUWI - ULSI 
oSuoD np anbyvisowad~ onbiyqnday 

sısuoun.tauds ofng 

STI 660 Pls ECO O81 SS7c SOF IVATO OSS 6% 891 VL IS REN COC 569 I STE} 


o8u09 np anbnelowoq anbiyqnday 
SISUOUMLOUDI ofng 


CEA £O'T 909 DONC 
ICI SL'0 ART VG Se 
8C I 8580 Ssor S6TO S 


SO RESO TIE EE 69 OV VLI OL Ir SO) LUNLUTXEIN 
ANOS CHA SES LAN Ze ivis L'6 rs ce Ll Lol DES LUNUITUT]A 
DL YOGI, ~ELViO 29S SCI 8967 Oria SOT (09 SELE Cel ISVs CTRO S 17 SOJUUI - ULSI 


QUO] LAIAIS BLIOQTT BURN) 
SISUIUNIIUDI ofng 


an 
nun 
am 


Pay PIN ISU] JOJOUPIP aappweıp SUA]  UIPIN o. Aso u 
HA/NS Melo Md/MH ‘Id/L Td/MH Md/Id MH/L TH/MH MH/NS 91418049) plowavg plowarg Ag wnurduidy, peop  PrOH  -Mous 


MILLS TANDY & JEAN-LUC PERRET 


398 


IVI 
st 
0ETO'O 
ICE I 
IVI 
ICI 
781000 
IOE I 


BEA 
STI 
cEIO'0 
Loc | 


LyT 
LOI 
r0r0°0 
ESCI 


9WI 
tcl 
TEcO'O 
€67 | 
dal 
OCT 
t1c0'0 
S8C I 


HA/NS 


ell 
tL0 
6cr0'0 
9760 
PAT 
890 
SLV0'0 
198°0 


£9'0 
8€ 0 
87c0'0 
SLr 0 


t9'0 
1E'O 
O££0'0 
8ct 0 


98 0 
8r 0 
IrE0'0 
£89'0 
960 
eco 
PILO'O 
60L'0 


Md/L 


18/S 
96€ 
61610 
8867 
STS 
GLE 
SYST'0 
Levy 


8S 
SEE 
76010 
0S6 € 


LES 
[TE 
DATA 
OLG'E 


IT'S 
LESE 
81610 
SLTY 
Ors 
vel 
EILEO 
LIGE 


Md/MH 


0€°0 
TA) 
8L00°0 
9LTO 
ECO 
veo 
L800°0 
8L70 


dat 
VARO) 
SLOO'0 
8910 


810 
cI0 
5900°0 
9710 


9£'0 
8c 0 
6L00°0 
SIEO 


Td/L 


691 
611 
ÿIS0'0 
por 


95] 
OT] 
tLv0'0 
6CE I 


65T 
PSI 
£660°0 
r06'1 


91T 
LAS 
r0L0'0 
OSL'I 


Id/MH 


(1087 
OST 
69ÿT'0 
MCE 
coe 
OST 
ISTIO 
880° 


LVE 
GET 
99710 
C86'C 


ITE 
781 
cOrl 0 
06CT 
167 
9L'0 
8SSITO 
LyTT 


Md/'Id 


0c 0 
910 
1€00°0 
S810 
tc 0 
LIO 
c800°0 
€610 


STO 
60°0 
L900°0 
ICI 


cI0 
60°0 
LE00'0 
OLIO 


810 
vIO 
6t700°0 
0910 
ITO 
STO 
1900°0 
1810 


MH/L 


0ÿc0'0 
0791 
LS 
9 1 
LSCO'0 
8ILT 


(0404 
9L'I 
8590°0 
L80°C 


[OT 
ELI 
SO£O'0 
SO8 I 
S6 I 
91 
15900 
EPL'I 


“TH/MH 


CLC 
861 
Lt90°0 
dd are 


CLIC 
OCT 
tcr0 0 
[IST 


MH/NS -2[{S01N ployed 


976 
LESS 
6b I 
80°88 


6 SCI 
098 


[OSE 
69°001 


geh 
OLE 
878'0 
r9°0r 
8:05 
Str 
€OL'0 
IZ Lp 


[904 


Or 
rol 
cIEO 
cL Il 


Pcl 
£'6 
IEIO 
Oc el 


c9 
6€ 
BETO 
65 
€91 
Ot 
LOT'T 
ts 9 


pia 


0°8E 
IT 
0960 
90'£E 


[op 
E TE 
csc I 
LL'8E 


sel 
08 
6850 
IE OI 
PSI 
OO! 


pos 0 
8S CI 


YU] 19)ALULIP_I19)QUIEIP 
ployeieg 


86 
88 
OCTO 
tc 6 
LOI 
T8 
€600 
€96 


CEI 
vs 
€550 
LYOI 


gel 
OLI 
SLTO 
CLI 


L'9 
es 
SCI O0 
065 
89 
CS 
SITO 
IT9 


9910 
16'S 
v8 
LS 
IS€0 
689 


99 
LY 
pOT 0 
IS’S 


IDE 
VT 
lEILO 
60'¢ 
9 
GE 
OST0 
SO UE 


SIC 
8 LI 
Src'0 
AA 
9°CT 
sl 
£OS'O 
S9'0C 


SIT 
£'6I 
SI80 


OC TC 


80€ 
LIT 
[C6 0 
6977 


OI 
68 
LSTO 
IE OI 
8SI 
£'OI 
£95°0 
IL'TI 


YU] 


akq wnuedwä] pray 


S'SE 
8/87 
0790 
LS'TE 
[8€ 
LACTA 
SS6°0 
Oÿ'SE 


8 96 
Liv 
O8e I 
cI'9y 


c'88 
LOL 
9811 
ells 
0'86 
9'YL 
IST 
16°68 


WNLUTXe 
wnwmu 


JOG PIPPUEJS 
0€ sojew - LOW 


wnwmxeN 

WINUITUI]A] 

JOLY parpuris 

OE SAW] - uray 


RLIISIN ‘OYSOWOgsa 


SLI 
0'801 
108° 1 
ST+II 


SLIDINSAL Ofng 


wnumxeN 

WUNLULUT AY 

IOLA parpuris 

Ol sopeu - ROW 


0800) np anbne.mowag anbrqndoy 


195 
OLY 
168°0 
LO'TS 


CIC 
l'O 
9750 
Ot 6l 
OT 
661 
A 
98 IC 


‘pia 
pray 


OLYVI 
VLII 
cele 
tC 8Cl 


c9°09 


afA}soin 
-jnous 


UMLWIXENN 
UUNUILUT]A 


JOLIG pICpureIs 
OI Safe] - uvoW 


08007 np anbyesowaq anbi[qndoY 77 unosgwed 


s1upIprosadns ofng 


WINXP IA] 

LUNLULUT IA 

JOM prepurys 

Ol sapeu - ROW 


wnwmxe N 
wunwmurg] 


JOLY paepurys 
OI so[eulay - URAN 


BLIIGIT ‘PIAOIUONN 
snıppnapu Ofng 


u 


399 


THE BUFO TUBEROSUS SPECIES GROUP 


Oro 861 STE 


T'GiOr CET CRC rsr 67 oc 


LST 001 867 89 Ir vol LIT LUNUIXETN 
ECI 990 vor GLU AT 90% VIO 881 08° DSS. OW, 66 rs LT 58 8 LI SLY CON CUTOTTN 
I0£00 LOE0O 9ÿ600 CrIOO 9L500 ITOIO 8900 0 S870°0 EIEOO 6860 £EOTO 8060 8510 ITIO colo LEEO C860 Jog PIEPUEIS 
ECO CS D 2579: STE DS SLT SOIT = 80 FEINE 969: 7 75508 0 eel 9 85€ 6€6 8L6l lees OT SOTEUI - UE 
IST 180 ISt TEO col VSG 10 Spia 16°C VALUE SORE CEI 99 cr sol I Ic 165 UINUUT XB IAT 
STI Lr'0 ILT LEO 8171 patta SIOE 57655 [ve LY 66 SS UE 18 vst 00S UUNUITUI]A 
LITOO  SLEO0 OS8TO 08000 99100 0£01°0 #S00°0 67900 8TE0°0 BCCI IETO 6€£0 6600 8110 00€0 I9£0 L060 10119 pIepuels 
89€ SSIO GILE EOEO 8721 6507 9LT0 PITT SELT LOO SoS SII LO9 LSet pré 8c0c css OT S®EUI9J - USM 
[eJauas 
luojuad ofng 
ESI 180 L8'V LEO] 2821 80€ 610 01T 667 89 BL LOT 88 sr ULI TLT 6 SL WINUNXETN 
LETI sso see ICO PTE ICT SIO: 672697 ST CS SEI 9 SE OI 6 IT 509 LUNUITUTIA 
8LC0°0 1900 I9ETO EOI0O ZLP0'0 StL0'0 070000 LLSO'O TST0°0 ILET 0670 1890 £EO£O 8II0 9950 OLSO 6071 JOUA P-EPURIS 
eIvt COLO Ir e970 PST SLOT OLIO 691 1087 IPOS €79 8591 SL CE ET TETE EC CEE OI sojelu - UESM 
LS’ I 88 0 87 I£O TI coe 070 SET E0°€ OCS v9 8 SI 970 gr L'ST 977 9EL LUNUNXETN 
cel L9'0 LESE LEQ: Es VEGAS NO EVS: 0er rs 8 PI 69 er GTI VEC 9°L9 NUN 
csy I TOLION NEST 160 S891 Sco? 9810 O1ZT [98'S -L9)tp> SBS \ SoS Lek thy Gyl mela 0169 Ç SOTEWIS] - UESW 
peu) ‘euauwelp N 
so.ax ofng 
peau UIPIM = YASUI] JOJOLUPIP JAJAUUPIP Sus, pin  ofAlsomm u 
HIVY/NS Md/L  Md/MH AId/L Id/MH Md/Id MH/L TH/MH MH/NS AIS, plowed plowwirg  oAg wnuedw A] peoH pes nous 


400 MILLS TANDY & JEAN-LUC PERRET 


TABLE 2. Apparent diagnostic differences between linear morphological characters of adult 
females of Bufo amieti and species of Bufo from west and central Africa and those all species of 
the B. regularis Comples. * species known from west of the Dahomey Gap. These comparisons 
are tentative because of the small sample size for B. amieti (n=2). A difference (+) indicates 
that ranges of variation do not overlap. These comparisons cannot be tested for statistical 
significance because of the small sample size for B. amieti and a few of the other species. For 
data on other species, see Table 7 in Largen er al. (1978), Table 12 in Tandy et al. (1982) and 
Tables 1, 9 and 16 in Tandy & Feener (1985). 


© © 
= =) > 2 7 
| ed i | 
Species DE à RME an 2 32 
Sex py yar AE NS EN ac ON ec ec CE | ws SAS 
Females 
B. tuberosus er E ati + + + 8 9 
B. gracilipes + + + + ++ + 7 10 
B. villiersi + + + + + + + + + + + # 12 5 
B. regularis* + + + + + + + + + + + 11 6 
BaSUDENCIILONIS mcr haan ASSE EER ts + + + + + + 14 3 
B. garmani + + + + + + + + + + + 11 6 
B. latifrons + + + + + + E EE nl 6 
B. togoensis* IB. Sp) geo! ae Dar Ar + + + + + 112 5 
B. brauni + + + + + + + + + + + + + + 14 3 
B. gutturalis Fall Epto. Peso = + + ee a in 6 
B. kisoloensis + + + + + + + + SPE + 12 5 
B. poweri È SP Wt. Apa enon ae AR. ara dr ESEL u} 14 3 
B. rangeri Ai ap, Gp SP Ge ir + + © 10 7 
B. camerunensis* + + + + + + + + + E LS tk 13 4 
B. kerinyagae + + + + + + + qi) 
B. xeros* A A e NUE A + PAE ar a ae 12 5 
B. asmarae + + + + + + - + + + + + 12 5 
B. maculatus* papi CES Ea a - + + + + + + 13 4 
B. danielae* + + + + + - + + + 9 8 
B. blanfordi + 4 + IG: > 12 
B. langanoensis + + + + + + O Hi 
B. turkanae + + + + + + + + + + + + + 13 4 
B. perreti CO PAU A SEA IO. - + Su tah fe hd Sb 13 4 
B. steindachneri + + Sey CE eos 8 9 
B. pentoni* 
- Senegal + + + + + Si + + + 


+ 
© 
N 


sharp; horizontal diameter of eye greater than length of snout; tympanum distinct, 
vertically oval, its horizontal diameter 43% that of the eye, 12% that of head width, 
tympanum diameter (T) 1.8mm, eye diameter (E) 4.Imm; parotoid glands enormous 
in relation to body size, globose, 1.6 times longer than wide, (PL/PW 1.60), narrowly 
separated from the eye, the anterior edges lying between the anterior and posterior 
borders of the tympanum, parotoid length (PL) 9.6mm, 26% of snout-urostyle length, 
parotoid width (PW) 6.0mm, 63% of parotoid length. 

First finger longer than second, second longer than fourth; fingers lacking 
marginal folds; subarticular tubercles large, undivided; palm with numerous tubercles 
lacking spinules; inner metacarpal tubercle moderately developed but much smaller 
than outer metacarpal tubercle and lacking spinules; lacking any series of distinct 
warts on the posterior surface of the forearm; toes moderately webbed; toe IV with 
almost four phalanges free of webbing on both the inner and outer margins; most toes 


401 


THE BUFO TUBEROSUS SPECIES GROUP 


OC 0 0 OC 0 a; NESM E I 12 S (0) Jussgqe 07 OI Ol 
[esouas - ..Juojuad ofng 
IE I 0 TE I + + da GI I 0 0 Juosard ce cl OC 
'0.19Xx Ofng 
pol 0 8L L6 S8 ur a (D) - 91 I 0 0 Juosard cl cs LCI 
1D/n921 ofng 
€ 0 SI cl 8 ne ne st: 0 0 0 OI juasqe OC OI OI 


0809 np anbne.mowpg anbrqnday 2 unoawre) - ,s11D1po1odns ofng 


SI OC LE LS 9I pi HEM + € OC SI 0 juasaid PL ce Ir 
„snippnapu Ofng 
b I t r SO Ù + 9 0 0 O 1uosad 6 € 9 
030L - :$1$U2080] OI 
TEE 9 SI IG I ap + + CC 0 0 0 juasoid 97 v CC 
[eloyenog vourny ‘TUNJA OLY 7 UNOIUPI - SU01/1D) ofng 
6 8 LI ve = See + (T) + LI 0 0 0 juasoid SE SI LI 
JUOIT HIS ‘OSuOD np anbiggnday ‘ELIOSIN ‘eLaqry] ‘aauiny “TeLIoyendg-Iung OLY ‘PUEUO ‘AIIOAT,P 9109 “UNOIBWILD - „SIsuounıaums ofng 
0 0 ST II PI de yeom : 14 L IT 0 juasoid SC € CC 
AMOAT,P AOD ‘PSESOUOIA - ..avjaiupp ofng 
8 0 0 8 0 - LACM = S 0 0 0 Juasqu OI r 9 ısaona ofng 
0 € LI GI I = yeom È 8 I I (0) juasoid 07 OI OI 
sadij19043 ofng 
Cc 0 9€ IC L = sti (DE G € I C juosqe 8€ DE 8 
o8uo9 np anbne.mowog anbrqnday ‘Ron vourny ‘unos - susosagny ofng 
0 0 G I I 3 Hr È & é é à é c G 0 
DOA], P 9109 - „HonuD ofng 
10[09 
10[09 Ie Mm ur 
= (+) + Jussgqe juasaid [ee ig Ho] JUS yuasqe wsıydıowıp  J[PJOL SPpewoJ sopew 
WA SM AL dd JM ‘JEU ur sguruodo ses [PIOA |enxos u somads 


‘SIBA O]UI U9YO1Q SIWNDWOS P[OJ [ESP] (€) ÂAJUO SIEWOJF Ur ANUS} UO pal (7) ‘10[09 [eJoWwe} pal aavy squawndads May AIDA (1) ‘(7861) 
‘yp Ja Apueg ur TI [QLL pue (8L61) ‘Te 19 USI] ur / [qu] 298 ‘satoads J9Y]O UO BEP 10, "Fummow [LUDA - WA UNS [esa] 10 [PSIOP uo 
sjods aNYM - SM ‘PIOJ [PS2] - LL ‘SPUejS projosed younsip - JA ‘UPS [PIOWIP] UO 10]09 Pol - HM Uns [BLOW 9] UO 10109 pal - IN den Aowoyra 
ay} JO ISOM WIOIJ UMOUY S919dS .. “BOLIPy [ENU99 pur Jsam WO ofng JO Soloods ur SIQIOVIEYI [EIITO[OYCIOWI IVAUT]UOU JO UONeLMA ‘€ TTAVL 


402 MILLS TANDY & JEAN-LUC PERRET 


with only a slight margin of webbing at the base; webbing with small melanized 
spinules on the dorsal surface; tubercles of soles smaller than those of palms; inner 
and outer metatarsal tubercles prominent and suited for burrowing, the inner larger 
and more raised than the outer. No tarsal fold. Urostyle-heel length (UH) 26.5mm. 
Dorsum with numerous conical and rounded warts, each with one or more large 
spinules and some smaller spinules; spinules extending over entire dorsum, including 
surfaces of parotoid glands, but less numerous anterior to eyes. Venter with somewhat 
smaller warts with spinules similar to those of the dorsum. Warts at rictus partially 
fused and covered with numerous small spinules. 

Ovaries with immature eggs. The specimen was apparently collected outside of 
the breeding season. 

Color (in alcohol) disruptively patterned light and dark brown; Dorsal back- 
ground color light brown; four pairs of bilateral reticulate dark brown markings. No 
B. regularis-like white spots on dorsum; No vertebral line; Parotoids and rictal warts 
same color as dorsum; Dark melanized spinules prominent against lighter back- 
ground. Lower margin of orbit and part of area beneath eye cream. Upper surfaces of 
limbs with well defined brown cross bars sometimes outlined in darker brown; 
posterior femoral integument light brown. Venter markedly reticulately mottled 
brown and cream. See Figs. | - 2. 

Paratype: The paratype female is similar to the holotype (Table 1). It is slightly 
smaller- SU 35.9mm, but is gravid and thus indicates that the breeding season 
includes December near the beginning of the long dry season. Coloration is like the 
holotype except overall a somewhat darker brown, and it has a few B. regularis-like 
white spots on the dorsum. See Figs. 3-4. 


GEOGRAPHIC DISTRIBUTION AND ECOLOGY 


B. amieti is known from only two localities in lowland rainforest of western 
Cöte-d’Ivoire (Fig. 5). Statistics on the altitude, rainfall and temperature of these 
localities are given in the diagnosis and in Table 4. 

The small number of known localities for B. amieti precludes statistical 
comparisons of ecological variables with other species. Data in Table 4 and 5 indicate 
some differences and similarities. B. amieti seems to differ in one or more ecological 
characteristics from eighteen of the 25 species with which it has been compared 
(Table 5), and it is unlikely to be sympatric with any of those. Of the remaining seven 
species, four, B. regularis, B. superciliaris, B. camerunensis and B. maculatus, are 
known from west Africa west of the Dahomey Gap, but B. amieti has not yet been 
found syntopic with any other bufonid species. Bufo cristiglans Inger & Menzies, 
1961, is considered a synonym of B. camerunensis Parker, 1936. 

If B. amieti is restricted to relatively undisturbed lowland rainforest, it is 
unlikely to occur in the same local areas as B. regularis or B. maculatus. But such a 
preference for high forest would make it more likely to occur with B. superciliaris or 
B. camerunensis. If B. amieti does occur locally with these species, it would probably 
avoid them because the small size of B. amieti would make it likely food fare for the 
other two species. 


403 


UP 


EROSUS SPECIES GRO 


THE BUFO TUB 


‘SASO12qN] ‘ = Soi) uodo ‘u ‘ds yonum 'g 


= SSOI9 YIM SIMI SHSO12QM 'g pue 1 


onup ofng JO UONNALISIP ILyde13093 UMOUY ‘G ‘DIH 


0 M 0% 


404 MILLS TANDY & JEAN-LUC PERRET 


TABLE 4. Ecological characteristics of Bufo amieti and other species of Bufo that are known to 
occur in west or central Africa. Accuracies and sources of these data vary. Some figures are 
based on surveys or meteorological stations near the locality. Others were interpolated from 
map contours, isotherms or isohyets. One decimal point given does not necessarily imply such 
accuracy of measurement but rather that these were the figures used to compute the statistics. 
Individual values for some altitudes may be in error by 50m or more. Most basic data are from 
Bartholomew (1985), Knoch & Schulze (1956), NIMA (1999), Wernstedt (1959, 1972) or 
Survey of Kenya (1962). 


n Elevation Mean Mean 


meters Annual Annual 
Temp oC Rainfall cm 
Bufo amieti 
Cöte d’Ivoire 
Grabazouo 150 23.9 161.1 
Tai 123 24 186.3 
Mean 2 136.5 23.95 1737 


Bufo tuberosus 


Cameroun, Gabon, Guinea Ecuatorial, République Centrafricaine, République du Congo, République Démo- 
cratique du Congo 


Mean 412.8 23.69 224.87 
Standard Error 46.63 0.316 20.472 
Minimum 10 22 150 
Maximum 1266 26.4 411.8 
n 48 14 15 
Bufo danielae 
Côte d'Ivoire 

Monogaga ] 10 25 150 


Bufo camerunensis 


Cameroun, Gabon, Nigeria, République du Congo & République Démocratique du Congo 


Mean 10 526.9 23.1 184.4 
Standard Error 95.49 0.49 13.92 
Minimum 100 20 140 
Maximum 1066.8 24.7 290 
Bufo togoensis 

Togo 3 447,3 22.8 141 
Minimum 350 22.8 139.7 
Maximum 589.8 22.8 142.3 
n 3 I 2 


Bufo latifrons 
Cameroun & Guinea Ecuatorial 


Mean 5 549.7 24.1 250.9 
Minimum 50 22.4 145.3 
Maximum 876.9 232 350 


Bufo maculatus 


Angola, Bissao, Cameroun, Ethiopia, Kenya, Liberia, Mozambique, Republique Centrafricaine, South Africa, 
Zimbabwe 


Mean 10 480.5 231 163.4 
Standard Error 206.62 1.09 44.4 
Minimum 19.5 26.7 491.2 


Bufo superciliaris 
Cameroun, Ghana, Republique Democratique du Congo 


Mean 10 561.1 22.9 193.9 
Standard Error 113.59 0.55 123,112 
Minimum 50 20 150 


Maximum 1066.8 26 260 


Bufo regularis 


Algeria, Angola, Cameroun, Egypt, Ethiopia, Gabon, Kenya, 


République Démocratique du Congo, Senegal, Sudan, Uganda 


THE BUFO TUBEROSUS SPECIES GROUP 


405 


Elevation Mean Mean 
meters Annual Annual 
Temp oC Rainfall cm 


Libya, Nigeria, République Centrafricaine, 


Mean 793.6 2117, 102.67 
Standard Error 81.16 0.453 8.373 
Minimum 7 14 0.3 
Maximum 2500 29.6 280 
Bufo xeros 

Chad, Ethiopia, Kenya, Uganda 

Mean 802.6 23.4 57.8 
Standard Error 164.72 1.41 7.8 
Minimum 54.9 16 29.9 
Maximum 1676.4 29.2 100 
Bufo pentoni 

Haute Volta, Senegal, Sudan 

Mean 131.5 DIET; 40.8 
Minimum 5 25 14.9 
Maximum 450 29 70 
Bufo gracilipes 

Cameroun, Gabon, Guinea Ecuatorial, Republique du Congo 

Mean 310.81 2392 214 
Standard Error 97.57163 0.345704 19.95348 
Minimum 10 23 166.4 
Maximum 780 23] 350 
Bufo villiersi 

Cameroun 

Mean 1853.5 23.38 263.62 
Minimum 1398 22.0 187.7 
Maximum 2500 25.0 305.0 


B. xeros and B. pentoni occur in much drier habitats in the northern portion of 
West Africa. 

B. danielae is known from only two coastal sites in Cöte-d’Ivoire (Perret, 
1977) which are at lower elevation and appear to have somewhat warmer and drier 
climates than do the known localities of B. amieti. 

B. togoensis is known from only three localities that are at higher elevations 
with somewhat cooler temperatures and lower rainfall than the known sites for B. 
amieti. Tandy (1972) considered B. togoensis to be a synonym of B. latifrons. Addi- 
tional recent collections indicate that B. togoensis is a distinct species resembling 
B. latifrons in the size and proportions of its tympani and 

B. camerunensis in the structure of its parotoid glands. 

The ecological features of B. tuberosus localities in central Africa do not 
appear to differ from B. amieti sites. 

The similarities of both morphology and ecology indicate a close relationship 
between B. amieti and B. tuberosus. Apparently B. amieti is a western isolate derived 
from a common ancestor of the two species. This pattern is similar to that of several 
pairs of lowland rainforest hyperoliid treefrog species east and west of the V-Baoulé 


406 MILLS TANDY & JEAN-LUC PERRET 


TABLE 5. Apparent differences and similarities between the ecological characteristics of Bufo 
amieti and species of Bufo from west or central Africa and those of all species of the B. 
regularis Complex. * species known from west of the Dahomey Gap. These comparisons are 
tentative because of the small number of known localities for B. amieti (n=2). A difference (+) 
indicates that ranges of variation do not overlap. These comparisons cannot be tested for statis- 
tical significance because of the small sample size for B. amieti and a few of the other species. 
(1) - species for which number of localities is less than 10. For data on other species, see Tables 
3 in Tandy et al. (1976) and Tandy er al. (1982), Table 8 in Tandy er al. (1985), and Table 13 
in Tandy & Feener (1985). 


Species compared Elevation Mean Annual Mean Annual Different Similar 


Temperature Rainfall 
B. tuberosus = & 0 3 
B. gracilipes - = 2 0 3 
B. villiersi (1) + È in 2 1 
B. regularis* - if 0 3 
B. superciliaris* 0 3 
B. garmani = = de I 2 
B. latifrons (1) - È 0 3 
B. togoensis*(1) + + fe 3 0 
B. brauni (1) + di St 3 0 
B. gutturalis - = At 1 2 
B. kisoloensis + + 2 1 
B. poweri + + fe 3 0 
B. rangeri = er | 2 
B. camerunensis* - 0 3 
B. kerinyagae chi + È 2 1 
B. xeros* An I 2 
B. asmarae + + DI 1 
B. maculatus* = A 0 3 
B. danielae* (1) + + ane 3 0 
B. blanfordi + = ae 2 1 
B. langanoensis (1) + + N 3 0 
B. turkanae (1) + SÈ n 3 0 
B. perreti (1) + + È DI 1 
B. steindachneri 2 i 1 2 
B. pentoni* - Senegal (1) - + + 2 1 


ecological break of Cöte-d’Ivoire as described by Schigtz (1967) for Hyperolius 
sylvaticus ivoiriensis vs H. s. sylvaticus, Leptopelis macrotis vs L. rufus (erroneously 
as “L. palmatus” (Perret, 1973)) and Leptopelis occidentalis vs L. boulengeri. The 
initial evolutionary barrier to eastern and western isolates probably is related to 
changes in Africa’s climate during geologic time and particularly to interpluvial 
periods when the rainforest blocks were probably most isolated. 


Bufo tuberosus Günther, 1858 


Bufo tuberosus Ginther, 1858: 60, pl. 3. fig. C, 140. Peters, 1875: 202. Boulenger, 
1880: 546-547, 564, 572-573; 1882: 304. Rochebrune, 1884: 20, 47-49, Pl. IV, Fig. 2. 
Boulenger, 1887: 565. Bocage, 1895: 15, 272. Werner, 1898: 202. Mocquard, 1903: 215. 
Boulenger, 1900: 435; 1903: 62. Bocage, 1903: 45. Boulenger, 1906: 158. Johnston, 1908: 
950. Nieden, 1908: 509-510, 517. Muller, 1910: 625. Nieden, 1910: 66, 68, Fig 147. Barbour, 
1911: 135. Lampe, 1911: 220. Arldt, 1917: 123. Noble, 1922: 39. Nieden, 1923:104. Noble, 
1924: 167, 177-178, 311. De Witte, 1930: 241-242, 252; 1934: 167. Loveridge, 1936: 84. 
Parker, 1936: 155. Mertens, 1938:10; 1941: 277. Monard, 1951: 174. Curry Lindahl, 1956: 56. 
Perret & Mertens, 1957: 555, 557. Guibé & Lamotte, 1958: 243. Inger & Menzies, 1961: 594. 


THE BUFO TUBEROSUS SPECIES GROUP 407 


Mertens, 1965: 14.19. Oates, 1965:89. Perret, 1966: 309-311, 314-315. Amiet & Perret, 1969: 
120. Perret, 1971: 131. Perret & Amiet, 1971: 48, 50, 52-53. Amiet, 1973: 136; 1975: 46, 50, 
54, 56 (photo); 1976: 150-152 (sonagram); 1978:199. Frost, 1985:63. Amiet, 1989: 92, 95, 98- 
99 (PI. I - photo). Brauer, 1991:6, 159, 179. 


Bufo polycerus Werner, 1897: 211. Nieden, 1923: 105. 


CAMEROUN - Akak-Yemefek; 2°37'N 10°04'E, alt. 50m; 2 MHNG917.67; 15.V.1955; J.-L. 
Perret. Batouri; 4°35'N 14°24'E, alt. 655m; 6 BM1934.12.1.14; IV.1933; M. Merfield. Bibundi 
(Bibunde); 4°13'N 8°59'E, alt. 50m; (Lampe, 1911). Bipindi; 3°06'N 10°30'E, alt. 200m; 1 
specimen; Zenker. (Nieden, 1910). Campo; 2°22'N 9°48'E, alt. 10m; 2 MHNCF148; 1.1947; 
A. Monard. Ebonji; 4°06'N 9°24'E, alt. 50m; "cri de rivalité" sonagram; (Amiet, 1989, p. 151). 
Efangono (=?"Efayong"); 3°04'N 11°49'E, alt. 750m; 9 BM1909.7.9.14; 1908?; G. L. Bates. 
Efulen, Kribi; 2°47'N 10°32'E, alt. 500m; d CNHM3585; Date? ; G. L. Bates. 2 BM1904. 
10.26.28; 1903? ; G. L. Bates. 9 MHNB2482; 1906; Rosenberg. Ekekam III; 3°53'N 11°22'E, 
alt. 750m; 6 MHNG1 176.5; 10.X.1968. 1 specimen; VI.1968. 3 specimens; X.1968-1.1969. J. 
-L. Amiet. Foulassi; 2°59'N 11°58'E, alt. 650m; 2 d& MHNG1008.30-31; 2 MHNG917.66; 
IV.1953; 49 2 MHNG1008.27-29, 1020.47; 8 im. MHNG1008.31-37, 1020.48-49; 1960-61; J. 
-L. Perret. Gadji; 4°29'N 14°03'E, alt. 600m; ?BM1937.1.1.21; 1936?; M. Merfield. Idenao; 
4°16'N 8°56'E, alt. 100m; 22 £ BM1968.470, -.508; 12.VIII.1967; T. Struhsaker. Iemonkonn; 
3°54'N 11°58'E, alt. 700m; 1 specimen; I-II/1968. J.-L. Amiet. Kala; 3(51'N 11(22'E, alt. 350m; 
recording of mating call; 12.1.1972; 2 specimens; VII-X.1967; 2 specimens; 15.II-15.1V.1969. 
J. -L. Amiet. Kendonge (Kidonge), S. Bakundu Forest Reserve; 4°33'N 9°26'E, alt. 50m; d 
BM1969.1635; 2 BM1969.509; 1968?; S. Gartlan. Kribi (includes "Dja River" of MCZ 
record); 2°56'N 9°56'E, alt. 43m; 2 MCZ2660; 1908; G. H. Schwab. 39 2 UMMZ 35611; I- 
VI.1907; 38320; 1908; 56277; no date; G. H. Schwab. Kumba (=Johann-Albrechtshöhe); 
4°38'N 9°25'E, alt. 200m; 2 specimens; Conradt. Nieden, 1910). Limbe (Victoria); 4°01'N 
9°12'E, alt. 100m; 4 specimens ZMB?13915; Preuss. (Nieden, 1908, 1910). 32 km N Lolodorf; 
3°26'N 10°42'E, alt. 250m; d CAS103353 (TP3545); 21-22.III.1966; T. Papenfuss. Lomié 
District; 3°09'N 13°35'E, alt. 632m; 2 BM1937.12.1.54; 1936?; M. Merfield. Mabiogo; 
2°12'N 9°53 E, alt. 10m; 2 MHNCF88; 28.11.1947; A. Monard. Mbanlam; 3°17'N 9°54'E, alt. 
612m; 6 MHNG955.100; 27.IX.1957; J. -L. Perret. Momobelenga; 3°56'N 11°40'E, alt. 600m; 
3 sonagram of mating call; Date?; (Amiet, 1976, p.151). Mundame; 4°35'N, 9°31'E, alt. 100m; 
2 specimens; 1906; Rohde. (Müller, 1910). Ngam/Sangmelima; 2°47'N, 11°54'E, alt. 759m; 
3 22 MHNG955.97-99; 4.1.1957, 3.11.1957, 20.IX.1957; 1 specimen SMF52335; 17.VI. 
1955; J. -L. Perret. Nkolfep; 3°59'N 11°24'E, alt. 350m; 1 specimen; 11.1968; J. -L. Amiet. 
Nta Ali; 5°33'N 9°30'E, alt. 1266m; 3; 15.1V.1977; J. -L. Amiet (Amiet, 1978). Rio del Rey; 
4°44'N 8°37E, alt. 50m; 1886?; H. H. Johnston. (Bouleng-er, 1887). Sangmelima (includes 
"Zima Country" of BM records); 2°56'N 11°59'E, alt. 782m; 29 2 BM1906.5.28.150-151; 
1905?; G. L. Bates. d CNHM19918; 1899; A. I. Good. nr Ting (32 km S Akonolinga) Sta. 
141; 3°40'N 12°10'E, alt. 750m; record-ing of mating call, 4 MT.TC.CA123; 7.IV.1971. M. 
& J. Tandy. Yapoma (nr Douala); 4°02'N 9°49'E, alt. 50m; 9 ZMB27541; 1910; Schafer. No 
locality; ¢ ZMB8319; 1870-1880; A. Reichnow. 

GABON - Makokou; 0°34'N 12°52'E, alt. 516m; 6 MHNG2207.33 (MBG1117); 25.X.1964; 9 
MHNG2207.34 (MBG1277); 26.X1.1964; L. P. Knoepffler. Nzing-Ayong (=?Nzibelong); 
0°18'N, 9°50'E, alt. 50m; 26 d MHNG2207.31-32 (MBG129, 336); 2 MHNG2207.30 
(MBG128); 26.11.1964; L. P. Knoepffler. 

GUINEA ECUATORIAL - Bioko (Fernando Poo): Basilé (Bassilé), slopes of Pico de Basilé (Santa 
Isabel); 3°36'N 8°45'E, alt. 527m; 2 adults MB; 1894?; F. Newton. (Bocage, 1895, 1903-1905). 
Rio Iladyi Falls, south of Bioko (Moca); 3°20'N 8°40'E, alt. 914m; 2 BM1969.2447; 
18.1X.1964; University College Expedition. Luba (Bahia de San Carlos); 3°27'N 8°33'E, alt. 
400m; im. SMF?; 20.IX-24.X.1962; R. Mertens. (Mertens, 1965). Moca (Moka); 3°20'N 
8°40'E, alt. 914m; ZMB333999; XI.1939; H. Eidmann. (Mertens, 1941). Musola; 3°26'N 
8°37'E, alt. 500-800m; (Boulenger, 1905-06). San Antonio de Ureca; 3°16'N 8°39'E, alt. 10m; 
?ZFMK9291; 1.1963; M. Eisentraut. No specific locality, 9 BM1947.2.21.14(=BM1851- 
10.25.7) Holotype; 1850; Fraser? 


408 MILLS TANDY & JEAN-LUC PERRET 


Rio Muni: Cabo San Juan (Cape St. John); 1°15'N 9°30'E, alt. 50m; Martinez de la 
Escalera. (Boulenger, 1903). Macomo (Makomo); 1°43'N 9°49'E, alt. 350m. (Nieden, 1908). 
Rio Uoro-Rio Mbini (Rio Benito); 1°34'N 9°38'E, alt. 50m; 4 2 9 BM1900.2.17.109-112; 
1899; G. L. Bates. Rio Lana 2.4km N Evinayong; 1°39'N 10°39'E, alt. 750m; ?BM1965.1412; 
6.1X.1965; J. Oates. No locality; ZMB20035; Date?; Collector? 

REPUBLIQUE DU CONGO (Congo-Brazzaville) - Djamba; 0°05'S 15°43'E, alt. 350m; RG3159; 
XII.1924; H. Schouteden. (De Witte, 1930). 

REPUBLIQUE DEMOCRATIQUE DU CONGO (Belgian Congo; Zaire) - Bafwabaka; 2°10'N 27°50'E, 
alt. 750m; 2 AMNH8405; 1909-1915; Congo Expedition. Buta; 2°47'N 24°47'E, alt. 350m; 2 
specimens RG4402-03; 11.1925; G. F. De Witte. Epulu; 2°15'N 29°15'E, alt. 350m; 2 speci- 
mens; 9.1V.1952. K. Curry-Lindahl. (Curry-Lindahl, 1956). Kifuku (s/la Nawa), ‚alt. ; 
RG1595; VIII.1925; H. Schouteden. Medje; 2°25'N 27°18'E, alt. 750m; RG1054; IV-V.1910; 
Lang & Chapin. Medje; 2°25'N 27°30'E, alt. 750m; AMNH8404; IV-V.1910; Congo Expedi- 
tion. (Noble, 1924). Ngayu; 1°40'N 27°40'E, alt. 750m; AMNH8401-02; IV-V.1910; Congo 
Expedi-tion. (Noble, 1924). (Note: AMNH8403 is a d Bufo funereus.) 

REPUBLIQUE CENTRAFRICAINE - Tibili (=?"Tuburi, Narob"); 4°58'N 14°42'E, alt. 750m; 2BM 
1912.1.11.19; 1911?; P. A. Talbot. 


MORPHOLOGY 


Many aspects of the morphology of B. tuberosus have been described and 
illustrated in the literature. Illustrations include drawings in Giinther (1858), Roche- 
brune (1884) and Nieden (1910) and photographs of preserved material in Perret & 
Amiet (1971) and of living specimens in Perret (1966) and Amiet (1975, 1989). 


PLATE 1. Bufo tuberosus female, Ngam, Sangmelima, Cameroun. 


THE BUFO TUBEROSUS SPECIES GROUP 409 


TABLE 6. Some diagnostic differences between linear morphological characters of adults of 
Bufo tuberosus and species of Bufo from west and central Africa and those all species of the B. 
regularis Complex. * species known from west of the Dahomey Gap. A difference (+) indicates 
that ranges of variation do not overlap. Lack of a diagnostic difference does not indicate lack of 
a statistical difference. These comparisons have not been tested for statistical significance. For 
data on other species, see Table 7 in Largen er al. (1978), Table 12 in Tandy et al. (1982) and 
Tables 1, 9 and 16 in Tandy & Feener (1985). 


Species 3 = 
Sex = = EME = A = = 5 5, £ 
ARE, ORE: 

B. amieti* tp Bars ap En + + + 8 

B. gracilipes + + + ER Credit 6 1 

B. villiersi 0 17 

B. regularis* + 1 17 

B. superciliaris* + + + + + + + + + 9 8 

B. gammani + + 2: 

B. latifrons + + DIST; 

B. togoensis* + + DNS 

B. brauni + + + + + + + 8 9 

B. gutturalis + + + Sun HA 

B. kisoloensis + 1 16 

B. poweri + + + + + + 6 1 

B. rangeri + + + + + 5 12 

B. camerunensis* + 1 16 

B. kerinyagae + HTC + 4 13 

B. xeros* + + + + + + + if 10 

B. asmarae + + + + + + 6 1 

B. maculatus* + 1 16 

B. danielae* + + + + 4 13 

B. blanfordi + + DIS 

B. langanoensis + + 3 14 

B. turkanae + + + + + + 6 11 

B. perreti + + + + Ay WS) 

B. steindachneri + + + + + + + 7 10 

B. pentoni* - Senegal + + ZU 
Males 

B. amieti* a TAN ERNEST Ce oe a n) ? ? 

B. gracilipes + + 2 IS 

B. villiersi et + + 5) 12 

B. regularis* + + Zils 

B. superciliaris* + + + + + + + + + + + + 12 5 

B. garmani See MORRIS al SEHEN RE + + + 9 8 

B. latifrons + 1 16 

B. togoensis* + + 2 15 

B. brauni + + + + + + + + + 9 8 

B. gutturalis + He + ++ + + ++ 10 7 

B. kisoloensis + + + + + + Ol 

B. poweri + + + + + + + + + + + 11 6 

B. rangeri 2 A RE D INI SCH + + + + 10 7 

B. camerunensis* + + 2 15 

B. kerinyagae + + + + 5 12 

B. xeros* + + + + + + + fetal 

B. asmarae + + + + + 5 12 

B. naculatus* + 1 16 

B. danielae* + l 16 

B. blanfordi + 1 16 

B. langanoensis + + 3 14 

B. turkanae + + + 3 14 

B. perreti + + DOS 

B. setindachneri + + 3 14 

B. pentoni* - Senegal + + 2 AS 


410 MILLS TANDY & JEAN-LUC PERRET 


Günther’s description focused on the extreme wartiness of the skin, but did not 
give measurements or ratios of body parts. The sex of the holotype was not specified 
nor was its exact locality on the island of Bioko (Fernando Poo) (Günther, 1858). 
Boulenger later published a more detailed description with some measurements and 
verbal proportions, however he mistakenly identifed the holotype as a male and his 
measurements for that specimen differ from ones made more recently. 

The very prominent parotoids were noted by Boulenger (1880) and by sub- 
sequent authors such as Noble (1924), Perret (1966) and Perret & Amiet (1971). The 
data in Tables | and 6 demonstrate the markedly low ratio of parotoid length to width 
and also the very small size of the tympanum compared to parotoid width. The ratio 
of T/PW separates B. tuberosus diagnostically (ranges of variation do not overlap) 
from 18 (in females) or 21 (in males) of the 25 other African Bufo species compared. 
As summarized in Table 6, B. tuberosus differs diagnostically in at least one of these 
measurements or their ratios from all of the species compared except for females of 
B. villiersi. 

Boulenger (1900) mentioned a crimson vertebral line in some specimens from 
the Rio Uoro (Rio Benito), Rio Muni, Guinea Ecuatorial. That color has now faded in 
those specimens. 

Noble (1924) noted varıation in ventral coloration in his material from the 
eastern République Démocratique du Congo ranging from heavy to no dark mottling. 

Parker (1936b) gave ratios of body to foot length and interpreted this as 
demonstrating relatively short legs of B. tuberosus compared to other species. Perret 
(1966) gave ranges of variation of body and tibia lengths for each sex and ratios of 
parotoid gland length/width and body length/tibia length. These data do not appear to 
support Parker’s appraisal of relative leg length, although Perret did not give arith- 
metic means of his ratios (only ranges) nor did he test such for statistical significance. 
Although differences of means given in Table | have not been tested for significance, 
the data on ratios of snout-urostyle length/urostyle heel length do not support Parker’s 
judgement about leg length in this species. Those data indicate that B. tuberosus is 
one of the longer legged species. 

Perret (1966) noted the marked sexual dimorphism of body size in this species. 
Tables 1 and 7 illustrate this dimorphism in linear characters. All measurements are at 
least statistically smaller in males than females, and two are diagnostically different. 
Most ratios do not differ between the sexes, although females have statistically 
narrower heads relative to body size (SU/HW) and smaller tympani relative to paro- 
toid width (T/PW). 


TABLE 7. Sexual dimorphism of linear morphological characters in Bufo tuberosus. D - 
difference diagnostic, ranges of variation do not overlap. S - statistically significant difference 
P < 0.01. - no statistical difference 


Number of characters 

2 3 o 

= = 3 Bo rà don mer 

Boe 2 e a 2 È S BE Sl 2 ei) a à 2 
»D S$ SS SS SH S S CR Ne 7) 


THE BUFO TUBEROSUS SPECIES GROUP 411 


Perret (1966) stated that he had not found vocal sac openings in males except 
for one with a single opening on the left. Perret & Amiet (1971) made a similar 
statement. Table 3 shows numbers and locations of vocal sac openings in the eight 
adult males examined. Two lack openings. 


MATING CALL 


Basic physical structure. The mating call of Bufo tuberosus is a series of low 
pitched hooting sounds that lack the noisy quality of the calls of most African Bufo. 
They are also not as loud as those of many other species (Amiet, 1976). The physical 
structure of these calls is similar to that of species of the B. regularis, B. maculatus, B. 
blanfordi, B. funereus (except B. villiersi), and B. pentoni groups and also some 
species of the B. vertebralis Complex and B. angusticeps group (Tandy, 1972; Tandy 
& Keith, 1972; Tandy & Tandy, 1976). Each complex pulse train (series of hoots) 
contains a variable number of simple pulse trains repeated at a rate of about 1.6 per 
second in specimens from Cameroun (Table 8). Each simple pulse train contains 
about 17 passive pulses repeated at a rate of 87 per second (89.74/sec at about 24°C). 
The duration of simple pulse trains is about 0.2 second (.18 at 24°C). The low 
emphasized frequency is the dominant frequency at about 526Hz. Sonagrams show 
three harmonics above the low emphasized frequency, but these contain less than half 
the energy of the low emphasized frequency. These frequencies show a slight gradual 
rise at the ends of simple pulse trains. These data are based on calls of only two 
recorded specimens. See material studied for times and places. 

Amiet (1976) described and illustrated a sonagram of a call of a male B. 
tuberosus uttered while trying to clasp a plastic bag containing another male which 
Amiet termed a “cri de rivalité”. This call is structurally very similar to male release 
calls uttered by many Bufo species, and the context indicates that is what it was. It is 
very different from the mating call in structure and function. 


TABLE 8. Physical characteristics of the mating calls of Bufo tuberosus. PPR = passive pulse 
repetition rate; NPP/PT = number of passive pulses per simple pulse train; PT DUR = simple 
pulse train duration; PTR = simple pulse train repetition rate; LOW EMP = low emphasized 
frequency; DOM = dominant frequency; n = number of individuals per sample. 


N PPR NPP/PT PT DUR PTR LOW EMP DOM 


(/sec) (/sec) (/sec) (Hz) (Hz) 
Bufo tuberosus 
Kala, Cameroun 1 

84.80 18.60 0.230 1.30 547.0 547.0 
Sta. 141 nr Ting, 1 
Cameroun 89.74 15.96 0.178 1.92 505.0 505.0 


Circadian and seasonal calling behavior. In April, 1971, Tandy found a mixed 
chorus of Bufo gracilipes and B. tuberosus calling along a small stream at dusk in 
secondary forest near Ting, Cameroun. The B. gracilipes called in water holding onto 
living aquatic plants and vegetative debris in the middle of the stream. B. tuberosus 
called from very well-concealed sites among overhanging roots of forest vegetation 
beneath the undercut stream bank. Amiet (1973, 1976, 1989) was able to make 


412 MILLS TANDY & JEAN-LUC PERRET 


additional observations of this behavior. The above two species and B. villiersi are the 
only Camerounian Bufo known to typically call during daylight, and these species are 
atypical of most species in the genus. 

Amiet (1976) has found that B. tuberosus choruses are usually small, often less 
than a dozen individuals. He also observed one larger aggregation near YaoundE, 
Cameroun. 

In Cameroun, B. tuberosus breeds during the dry season and at the beginning 
of the rainy season fi principally January-March (Amiet, 1976). 

Comparisons with other species. The mating call of B. tuberosus is as distinc- 
tive as its morphology. Table 10 compares the acoustic structure to those of twenty 
African species including all species whose mating calls are known from west and 
central Africa and all members of the B. regularis Complex (Tandy & Tandy, 1976). 
See also data in Tandy et al., 1976; Largen et al., 1978; and Tandy et al., 1982 and 
1985. The call of B. tuberosus appears to differ diagnostically (ranges of variation do 
not overlap in data available) in at least 3 characters from all twenty species. This 
comparison must be considered tentative because of the small sample size (2) 
available for B. tuberosus, but the comparisons are probably valid because of the 
much larger data sets available for most of the other species. 

Only B. gracilipes has been observed calling at the same time and locality as 
B. tuberosus, although B. camerunensis has been observed breeding nearby. Other 
species of the central African rain forests that might breed in the same areas include 
B. latifrons, B. maculatus and B. superciliaris. The latter is thought to lack a mating 
call (Amiet, 1976). 

The call of B. tuberosus differs in five of six apparently homologous characters 
from B. gracilipes, B. villiersi and B. perreti and it also differs qualitatively from 
those species. The mating calls of the above three species are first order sequences of 
complex pulse trains whereas that of B. tuberosus 1s just one complex pulse train. 

The voice of B. camerunensis differs in all six call characters compared. The 
PPR is slower in B. camerunensis, the NPP/PT and PT DUR two to three times 
greater and PTR is one third to one fourth its rate in B. tuberosus. 

The. mating call of B. latifrons is different in all six characters. PT DUR in 2. 
tuberosus is about one fifth, PPR is more than three times faster and PTR is two to 
four times faster than in B. latifrons. 

The voice of B. maculatus differs in four characteristics. PPR and PT DUR are 
about twice, NPP/PT four times and PTR up to twice greater than in B. tuberosus. 

The call of B. regularis differs in three characters. PPR is less than a third, PT 
DUR is about four times greater and PTR less than one half those in B. tuberosus. 
These two species are not likely to be sympatric because of different habitat pre- 
ferences. 

None of the other species in Table 9 is likely to be sympatric with B. 
tuberosus. Their calls differ from the latter species in at least three characters. 


THE BUFO TUBEROSUS SPECIES GROUP 413 


TABLE 9. Apparent differences and similarities between the mating calls of Bufo tuberosus and 
19 species of Bufo from west or central Africa and all those known of species of the B. 
regularis Complex. * species known from west of the Dahomey Gap. “ species of central 
Africa. These comparisons are tentative because of the small number of specimens recorded for 
B. tuberosus (n=2). A difference (+) indicates that ranges of variation do not overlap. These 
comparisons cannot be tested for statistical significance because of the small sample size for B. 
tuberosus and a few of the other species. (1) - species for which number of specimens analyzed 
is less than 10. For data on other species, see Table 1 in Tandy et al. (1976), Table 2 in Largen 
et al. (1978), Table 1 in Tandy er al. (1982), and Table 14 in Tandy & Feener (1985). (2) 
Comparison is between homologous characters rather than those of the same grade of physical 
compexity (See Tandy & Tandy, 1976). 


Species compared PPR NPP/PT PTDUR PTR LOW EMP DOM Number of characters 
Different Similar 


+ (2) + (2) 
+ (2) + (2) 


B. gracilipes* + 
villiersi “(1) + 
. regularis** + 
garmani = 
latifrons (1) + 
brauni - 
gutturalis + 
kisoloensis (1) - 
poweri + 
rangeri + 
camerunensis*" +? 
+ 
+ 
+ 
+ 
+ 
+ 
+ 
+ 


lea 
"9 
+ 
So 


+++ + + 
+++++ +++ ++. 


kerinyagae (1) 

xeros* 

asmarae 

maculatus*" 
langanoensis (1) 
turkanae (1) 

perreti 
steindachneri”\(1) 

B. pentoni** - Senegal (1) 


+ 
~ 

+ 
~ 


++++++ +. 


matt vote ++ ++t+t+t+++ 
oe ++: 


+ 
Ic Ra rer 


Is Do Do iu Do Du iu Do Du io Do Du io bp in bu in I 
+ 


DAHNNADHNBRADHAHRWUADWWUY 
ON = ON = ND © © N D D & = © à & TT 


> 
+ +++ 
~ 


+ 


| 


GEOGRAPHIC DISTRIBUTION AND ECOLOGY 


Bufo tuberosus is known from fiftyone equatorial lowland rainforest localities 
in Cameroun, Guinea Ecuatorial (Bioko and Rio Muni), Gabon, République du 
Congo, République Centrafricaine and République Démocratique du Congo between 
latitudes 0°5’S and 5°14’N and longitudes 8°33’ and 29°15’E (Fig. 5). These are in an 
altitudinal range of 10 to 1266m. See Table 4 for climatic data. 

Ecological data in Tables 4 and 10 indicate that B. tuberosus localities differ 
diagnostically from those of fifteen of the 25 species compared. B. tuberosus is not 
likely to be sympatric with any of those. 

Among the remaining ten species, seven occur in central Africa, B. gracilipes, 
B. superciliaris, B. latifrons, B. camerunensis, B. maculatus and B. steindachneri. As 
noted above, B. tuberosus has been found syntopic only with B. gracilipes. But it is 
likely to occur within close proximity of all of these except B. steindachneri. The 
latter occurs primarily in marshy sites surrounded by arid savanna. 

B. tuberosus is known only from forested localities, so it is not so likely to 
occur with B. regularis or B. maculatus which often prefer more open habitats. 


414 MILLS TANDY & JEAN-LUC PERRET 


The other high forest species, B. gracilipes, B. superciliaris, B. latifrons and B. 
camerunensis are most likely to occur with B. tuberosus. 

As noted previously, the ecology of B. tuberosus sites is very much like that of 
the allopatric B. amieti. 


TABLE 10. Apparent differences and similarities between the ecological characteristics of Bufo 
tuberosus and species of Bufo from west or central Africa and those of all species of the B. 
regularis Complex. * species known from west of the Dahomey Gap. “ species known from 
central Africa. Some of these comparisons are tentative because of the small number of known 
localities for some species. A difference (+) indicates that ranges of variation do not overlap. 
These comparisons have not been tested for statistical significance. (1) - species for which 
number of localities is less than 10. For data on other species, see Tables 3 in Tandy et al. 
(1976) and Tandy er al. (1982), Table 8 in Tandy er al. (1985), and Table 13 in Tandy & 
Teener (1985). 


Species compared Elevation Mean Annual Mean Annual Different Similar 


Temperature Rainfall 
B- Amieti (1) = È = 0 3 
B. gracilipes * - 5 è 0 3 
B. villiersi * (1) + 3 I 2 
B. regularis *’ = 0 3 
B. superciliaris ** - = = 0 3 
B. garmani - È fu I 2 
B. latifrons * (1) = Z Le 0 3 
B. togoensis (1) = 2 TE I 2 
B. brauni (1) = N È I 2 
B. gutturalis = = Fe 1 2 
B. kisoloensis * = N x 2 1 
B. poweri = È + I 2 
B. rangeri = + = | 2 
B. camerunensis ** - = 0 3 
B. kerinyagae + + - 2 1 
B. xeros ** = 2 + I 2 
B. asmarae = = + 1 2 
B. maculatus ** = 2 È 0 3 
B. danielae * (1) - - = 0 3 
B. blanfordi = = st | 2 
B. langanoensis (1) = a a I 2 
B. turkanae (1) = ave SÈ 2 1 
B. perreti (1) = is r 0 3 
B. steindachneri” = x = 0 3 
= 2 


pentoni **- Senegal (1) - = 


ACKNOWLEDGEMENTS 


M. Tandy expresses his appreciation to the late W. F. Blair for providing funds 
via National Science Foundation Grant 5406X which supported his fieldwork. He also 
thanks his wife Judy for her continued encouragement and support. He thanks his 
former wife, J. Hooper for her assistance with field study. 

We thank J.-L. Amiet (Université Fédérale du Cameroun) for making available 
his recordings of Bufo gracilipes, B. latifrons and B. tuberosus. 

We are grateful to the following for providing preserved or living material 
from collections in their care: J.-L. Amiet; the late C. M. Bogert, R. Zweifel, C. W. 
Myers and G. Foley (New York); J. E. Böhlke, L. Keane and E. V. Malnate 


THE BUFO TUBEROSUS SPECIES GROUP 415 


(Philadelphia); W. Böhme (Bonn); A. G. C. Grandison and B. Clarke (London); J. 
Guibé, E. R. Brygoo, A. Dubois and J. J. Morere (Paris); M. S. Hoogmoed (Leiden); 
R. F. Inger and H. Marx (Chicago); M. Lamotte (Paris); A. E. Leviton, R. C. Drewes 
and T. Papenfuss (San Francisco); the late R. F. Martin, A. Anderson, D. Canatella 
(Austin); C. J. McCoy (Pittsburgh); The late R. Mertens and K. Klemmer (Frankfurt 
a. M.); J.-L. Perret and V. Mahnert (Genève); the late G. Peters, P. Beurton and 
R. Guenther (Berlin); the late J. A. Peters, R. Crombie, R. Heyer, F. McCullough and 
R. Zug (Washington); U. Rahm and C. Unternahrer (Bale); G. Sacarrao and M. 
Noqueira (Lisbon); E. Balletto, M. A. Cherchi, the late G. Scortecci and P. Versace 
(Genoa); the late R. C. Stebbins and D. B. Wake (Berkeley, California); the late 
D. W. Tinkle, R. A. Nussbaum and M. H. Eckhardt (Ann Arbor, Michigan); E. E. 
Williams, J. Rosado and P. Kerfoot (Cambridge, Massachusetts) and J. W. Wright 
(Los Angeles). 


LITERATURE CITED 


AMIET, J. -L. 1973. Notes faunistiques, éthologiques et écologiques sur quelques Amphibiens 
Anoures du Cameroun. Annales de la Faculté des Sciences du Cameroun 13:135-161. 

AMIET, J.-L. 1975. Ecologie et distribution des Amphibiens Anoures de la region de Nkong- 
samba (Cameroun). Annales de la Faculté des Sciences du Cameroun 20:33-107. 

AMIET, J.-L. 1976. Voix d'Amphibiens camerounais. V. - Bufonidae : genres Bufo, Werneria 
et Nectophryne. Annales de la Faculté des Sciences du Cameroun 21-22:139-157. 

AMIET, J.-L. 1978. Les Amphibiens Anoures de la région de Mamfé (Cameroun). Annales de 
la Faculté des Sciences du Cameroun 25:189-219. 

AMIET, J. -L. 1989. Quelques aspects de la biologie des Amphibiens Anoures du Cameroun. 
Année Biologique. Paris 28:73-136. 

AMIET, J. L. & PERRET, J. -L. 1969. Contributions a la faune de la région de Yaoundé 
(Cameroun) II. -Amphibiens Anoures. Annales de la Faculté des Sciences du Came- 
roun 3:117-137. 

ARLDT, T. 1917. Die Ausbreitung der Lurche. Archiv fiir Naturgeschichte 82:94-151. 

BARBOUR, T. 1911. Some West African Amphibians. Bulletin of the Museum of Comparative 
Zoology 54:127-139. 

BARTHOLOMEW, J. (ed.). 1985. The Times Atlas of the World. Times, London. 

BOCAGE, J. V. B, DU. 1867. Batraciens nouveaux de l'Afrique occidentale (Loanda et Ben- 
guella). Proceedings of the Zoological Society of London 843-846. 

BOCAGE, J. V. B, pu. 1895. Subsidios para a fauna da ilha de Fernäo do P6 — Vertebrados 

Terrestres. Jornal de Sciencias Mathematicas, Physicas e Naturaes, Lisboa 2 (13): 1-15. 

BOCAGE, J. V. B, pu. 1903. Contribution a la faune des quatre iles du Golfe de Guinée. Jornal 
de Sciencias Mathematicas, Physicas e Naturaes, Lisboa 7 (26):25-96. 

BOULENGER, G. A. 1880. On the Palaearctic and Aethiopian species of Bufo. Proceedings of 
the Zoological Society of London 640-646. 

BOULENGER, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of 
the British Museum. 2nd ed. London. 

BOULENGER, G. A. 1887. A list of the reptiles and batrachians collected by Mr. H. H. Johnston 
on the Rio del Rey, Cameroons District, West Africa. Proceedings of the Zoological 
Society of London 564-565. 

BOULENGER, G. A. 1900. A list of the batrachians and reptiles of the Gaboon (French Congo) 


with descriptions of new genera and species. Proceedings of the Zoological Society of 
London 433-456, pls. XX VII-XXXII. 


416 MILLS TANDY & JEAN-LUC PERRET 


BOULENGER, G. A. 1903. Batraciens de la Guinée espagnole. Memorias de la Real Sociedad 
Espanola de Historia Natural, Madrid 1(2):61-65. 

BOULENGER, G. A. 1906. Report on the batrachians collected by the late L. Fea in West Africa. 
Annali del Museo Civico di Storia Naturale Giocomo Doria, Genova (3)2:157-172. 

BRAUER, K. 1991. Kröten. Urania. Leipzig. 190 pp. 

CURRY-LINDAHL, K. 1956. Ecological studies on mammals, birds, reptiles, and amphibians in 
the eastern Belgian Congo. Annales du Musée Royal du Congo Belge. Tervuren (Bel- 
gique). Séries Sciences Zoologiques 42:1-78. 

Frost, D. R. (ed). 1985. Amphibian Species of the World. Allen Press and Association of 
Systematics Collections, Lawrence, Kansas. 

GuIBE, J. & M. Lamotte. 1958. La Réserve naturelle intégrale du Mont Nimba, XXI. Batra- 
ciens. Mémoires de l’Institut Français d'Afrique noire. Dakar 53:241-273. 

GUNTHER, A. 1858. Catalogue of the Batrachia Salientia in the collection of the British 
Museum. London. 

INGER, R. F. & Menzies, J. I. 1961. A new species of toad (Bufo) from Sierra Leone. Fieldiana 
Zoology, Chicago 39:589-594. 

JOHNSTON, H. 1908. George Grenfell and the Congo. Hutchinson, London. 

KnocH, K. & SCHULZE, A. 1956. Precipitation, temperature and sultriness in Africa. Hamburg: 
Falk-Verlag, 14 pp., 10 maps. 

LAMPE, E. 1911. Erster Nachtrag zum Katalog der Reptilien- und Amphibien -Sammlung des 
Naturhistorischen Museums der Stadt Wiesbaden. Jahrbuch des Nassauischen Vereins 
für Naturkunde. Weisbaden 64:137-236. 

LARGEN, M. J., TANDY, M. & TANDY, J. 1978. A new species of toad from the Rift Valley of 
Ethiopia, with observations on the other species of Bufo (Amphibia Anura Bufonidae) 
recorded from this country. Monitore Zoologico Italiano, N. S. Supplemento 10:1-41. 

LOVERIDGE, A. 1936. African reptiles and amphibians in the Field Museum of Natural History. 
Field Museum of Natural History, Zoological Series 22: 1-111. 

MERTENS, R. 1938. Herpetologische Ergebnisse einer Reise nach Kamerun. Abhandlungen der 

Senckenbergischen Naturforschenden Gesellschaft 442:1-52. 

MERTENS, R. 1941. Zur Kentnis der Herpetofauna von Fernando-Poo. Zoologischer Anzeiger 

135:275-281. 

MERTENS, R. 1965. Die Amphibien von Fernando Poo. Bonner Zoologische Beiträge 16:13-29. 

MOCQUARD, M. F. 1903. Notes Herpétologiques. Description d'espèces de la collection du 

Museum. Bulletin du Museum national d’Histoire Naturelle. Paris 9:209-221. 

MONARD, A. 1951. Résultats de la mission zoologique suisse au Cameroun. Mémoires de 
l’Institut Français d’Afrique noire, Centre du Cameroun 1:1-244. 

MÜLLER, L. 1910. Beiträge zur Herpetologie Kameruns. Abhandlungen der Königlich 
Bayerische Akademie der Wissenschaften. München 24:545-627. 

NIEDEN, F. 1908. Die Amphibienfauna von Kamerun. Mitteilungen aus dem Zoologischen 
Museum in Berlin 3:489-518. 

NIEDEN, F. 1910. Die Reptilien (ausser den Schlangen) und Amphibien. Die Fauna der deut- 
schen Kolonien. Reihe I. Kamerun. Berlin. 74 pp. 

NIEDEN, F. 1923. Anura I. Das Tierreich. Berlin and Leipzig. VII-XXII + 575 pp. 

NIMA (National Imagery and Mapping Agency). 1999. GEOnet Names Server. www.nima.mil 
National Imagery and Mapping Agency. Washington, D. C. 

NOBLE, G. K. 1922. The phylogeny of the Salientia. I. The osteology and the thigh musculature; 
their bearing on classification and phylogeny. Bulletin of the American Museum of 
Natural History 46:1-87. 

NOBLE, G. K. 1924. Contributions to the herpetology of the Belgian Congo based on the 
collection of the American Museum Congo Expedition, 1909-1915. Bulletin of the 
American Museum of Natural History 49:147-347. 


THE BUFO TUBEROSUS SPECIES GROUP 417 


OATES, J. 1965. Expedition to Fernando Po. Animals 7: 86-91. 

PARKER, H. W. 1936. The amphibians of the Mamfe Division, Cameroons. I. Zoogeography 
and systematics. Proceedings of the Zoological Society of London: 135-163. 

PERRET, J. -L. 1966. Les amphibiens du Cameroun. Zoologische Jahrbücher, Systematik. Jena 
93: 289-464. 

PERRET, J. -L. 1971. Redécouverte de Bufo villersi Angel. Revue de Zoologie et de Botanique 
Africaines 84:130-139. 

PERRET, J. -L. 1973. Leptopelis palmatus (Peters) et Leptopelis rufus (Reicheow) (Amphibia 
Salientia): deux especes distinctes. Annales de la Faculté des Sciences du Cameroun 
15-16: 81-90. 

PERRET, J.-L. 1977. Une nouvelle espèce de crapaud africain: Bufo danielae de Côte d'Ivoire. 
Revue suisse de Zoologie 84: 237-245. 

PERRET, J. -L. & AMIET, J. -L. 1971. Remarques sur les Bufo (Amphibiens Anoures) du 
Cameroun. Annales de la Faculté des Sciences du Cameroun 5: 47-55. 

PERRET, J. -L. & MERTENS, R. 1957. Etude d'une collection herpétologique faite au Cameroun 
de 1952 à 1955. Mémoires de l'Institut Français d’Afrique noire. Dakar 19: 548-601. 

PETERS, W. 1875. Uber die von Herrn Professor Dr. R. Buchholz in Westafrika gesammelten 
Amphibien. Monatsberichte der Königlichen Preussischen Akademie der Wissen- 
schaften. Berlin 1875: 196-212. 

ROCHEBRUNE, A. -T. 1884. Faune de la Sénégambie. Octave Doin. Paris. 

SCHIOTZ, A. 1967. The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei 
hauniensis 25: 1-346. 

SURVEY OF KENYA. 1962. Atlas of Kenya. Nairobi: Survey of Kenya, IX + 46 pp. 

TANDY, M. 1972. The evolution of African Bufo. Ph. D. dissertation, University of Texas, 
Austin. 

TANDY, M., BOGART, J. P., LARGEN, M. J. & FEENER, D. J. 1982. A tetraploid species of Bufo 
(Anura Bufonidae) from Ethiopia. Monitore Zoologico Italiano, N. S. Supplemento 17: 
1-79. 

TANDY, M., BOGART, J. P., LARGEN, M. J. & FEENER, D. J. 1985. Variation and evolution in 
Bufo kerinyagae Keith, Bufo regularis Reuss and Bufo asmarae Tandy et al. Monitore 
Zoologico Italiano, N. S. Supplemento 20: 211-267. 

TANDY, M. & FEENER, D. J. 1985. Geographic variation in species of the Bufo blanfordi Group 
(Amphibia: Bufonidae) and description of a new species (pp. 549-585). Zn: Schuch- 
mann K. -L. (ed.) Proceedings of the International Symposium on African Vertebrates. 
Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn. 

TANDY, M. & KEITH, R. 1972. African Bufo (pp. 119-170). In: Blair, W. F. (ed.). Evolution in 
the genus Bufo. University of Texas Press, Austin. VIII+459 pp. 

TANDY, M. & TANDY, J. 1976. Evolution of acoustic behavior of African Bufo. Zoologica 
Africana 11: 349-368. 

TANDY, M., TANDY, J., KEITH, R. & DurF-MACKAY, A. 1976. A new species of Bufo (Anura: 
Bufonidae) from Africa's dry savannas. The Pearce-Sellards Series. Austin 24:1-20. 

WERNER, F. 1897. Ueber einige neue oder seltene Reptilien und Frosche der zoologischen 
Sammlung des Staates in Miinchen. Sitzungsberichte der Mathematisch-Naturwissen- 
schaftlichen Klasse der Bayerischen Akademie der Wissenschaften. Miinchen 23: 
203-220. 

WERNER, F. 1898. Ueber Reptilien und Batrachier aus Togoland, Kamerun und Tunis I. 
Verhandlungen der Zooligisch-Botanischen Gesellschaft. Wien 48: 191-231. 

WERNSTEDT, F. L. 1959. World climatic data. Africa. Dept. of Geography, Pennsylvania State 
University, State College, Pennsylvania. 101 pp. 

WERNSTEDT, F. L. 1972. World climatic data. Lemont, Pennsylvania: Climatic Data Press, 
523 pp. 


418 MILLS TANDY & JEAN-LUC PERRET 


WITTE, G. -F. DE. 1930. Liste des batraciens du Congo Belge. I. Revue de Zoologie et de 


Botanique Africaines 19: 232-274. 
WITTE, G. -F. DE. 1934. Batraciens récoltés par le Dr. H. Schouteden et par M. G. -F. De Witte. 
Annales du Musée Royal du Congo Belge. Tervuren (Belgique). Séries Sciences 


Zoologiques 3: 53-188. 


REVUE SUISSE DE ZOOLOGIE 


Tome 107 — Fascicule 2 


LANG, Claude. Etat trophique du lac de Morat indiqué par le zoobenthos: 
tendance 1980-199 8 vee Ro a RR ANA nn UE 


AZPELICUETA, Maria de la Mercedes & José O. GARCIA. A new species of 
Astyanax (Characiformes, Characidae) from Uruguay river basin in 
Argentina, with remarks on hook presence in Characidae........... 


BALLERIO, Alberto. Revision of the genus Ebbrittoniella Martinez (Cole- 
optera+Scarabaeoıdea: Ceratocanthidae) Zr anna Need. 


LIENHARD, Charles. A new desert psocid from Namibia (Insecta: Psocoptera: 
Hroslidaeye ne en Te ee ER 


LEISTIKOW, Andreas. Terrestrial Isopoda from Guatemala and Mexico 
(Crustacea Oniscidea: Crinocheéta) > 


BURCKHARDT, Daniel & Mariängela GUAJARA. Euphalerus clitoriae sp. n., a 
new psyllid species from Clitoria fairchildiana (Fabaceae, Papiliono- 
ideae), and notes on other Euphalerus spp. (Hemiptera, Psylloidea). . . 


KLoss, Kerstin & Wolfgang PFEIFFER. Zur Biologie des “Eingeweide- 
fisches” Carapus acus (Brünnich, 1768) (Carapidae, Teleostei), mit 
Hinweisen auf eine nicht-parasitische Ernährung. ................. 


BEDOS, Anne & Louis DEHARVENG. Un nouveau Collembole Neanurinae du 
Sud du Vietnam, Blasconura batai sp. n., avec une clé des especes du 
genre(CollembolasNeanuridac) è Corr 


BAUR, Hannes & Felix AMIET. Die Leucospidae (Hymenoptera: Chalcido- 
idea) der Schweiz, mit einem Bestimmungsschliissel und Daten zu den 
CUFOpdischenvAnteMm ar o i Eee 


TANDY, Mills & Jean-Luc PERRET. The Bufo tuberosus species group with 
the description of a new species from the rainforest of Cöte-d’Ivoire. . . 


Pages 


233-243 


245-257 


259-275 


277-281 


283-323 


325-334 


335-349 


351-357 


359-388 


389-418 


REVUE SUISSE DE ZOOLOGIE 
Volume 107 — Number 2 


LANG, Claude. Trophic state of Lake Morat indicated by the zoobenthos: the 
NOS ORI OO So ae 120: ren te I EE 


AZPELICUETA, Maria de la Mercedes & José O. GARCIA. A new species of 
Astyanax (Characiformes, Characidae) from Uruguay river basin in 
Argentina, with remarks on hook presence in Characidae........... 


BALLERIO, Alberto. Revision of the genus Ebbrittoniella Martinez (Cole- 
optera: Scarabaeoidea:-Ceratocanthidae): EE a O 


LIENHARD, Charles. A new desert psocid from Namibia (Insecta: Psocoptera: 
Miro Silica) iver rs cc ein ord er ER ETS 


LEISTIKOW, Andreas. Terrestrial Isopoda from Guatemala and Mexico 
(Crustacea Oniscidea: Crinocheta)........ rn Sea eee 


BURCKHARDT, Daniel & Mariängela GUAJARA. Euphalerus clitoriae sp. n., a 
new psyllid species from Clitoria fairchildiana (Fabaceae, Papiliono- 
ideae), and notes on other Euphalerus spp. (Hemiptera, Psylloidea). . . 

Koss, Kerstin & Wolfgang PFEIFFER. On the biology of the pearlfish 
Carapus acus (Brünnich, 1768) (Carapidae, Teleostei), with indica- 
tions of amon-parasitic nutrition» +2 <= = eres ee RES 

BEDOS, Anne & Louis DEHARVENG. A new neanurine springtail from south- 
ern Vietnam, Blasconura batai sp. n., with a key to the species of the 
genus: (GollembolasNeanuridae)2.. 2.2. RARA de E 

BAUR, Hannes & Felix AMIET. The Leucospidae (Hymenoptera: Chalcido- 
idea) of Switzerland, with a key and data on the European species. . . . 

TANDY, Mills & Jean-Luc PERRET. The Bufo tuberosus species group with 
the description of a new species from the rainforest of Côte-d'Ivoire... 


Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX 


Pages 


233-243 


245-257 


259-275 


277-281 


283-323 


325-334 


335-349 


351-357 


359-388 


389-418 


PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE 


CATALOGUE DES INVERTEBRES DE LA SUISSE, N° 1-17 (1908-1926) ..... serie Fr. 285.— 


(prix des fascicules sur demande) 
REVUE DE PALÉOBIOEOGIE mL Nenn. Echange ou par fascicule Fr. 
LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris) 


COLLEMBOLENFAUNA EUROPAS 
von H. Gisin, 312 Seiten, 554 Abbildungen, 1960 (Nachdruck, 1984) ............. Fr. 


THE EUROPEAN PROTURA THEIR TAXONOMY, ECOLOGY AND 
DISTRIBUTION WITH KEYS FOR DETERMINATION 


are par fascicule Fr. 


by J. NOSEK, 346 pages, 111 figures in text, 1973.............................. Fr. 
CLASSIFICATION OF THE DIPLOPODA 
PARRIChAr dE HOREMAN; 237) pases) 1979 N oe ee Fr. 


LES OISEAUX NICHEURS DU CANTON DE GENEVE 
par P. GEROUDET, C. GUEX et M. MAIRE 
351 pages, nombreuses cartes et figures, 1983 ................................. Fr. 


CATALOGUE COMMENTÉ DES TYPES D'ECHINODERMES ACTUELS 
CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES, 
SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ 
A LA CONNAISSANCE DES ECHINODERMES ACTUELS 
par Michel JANGOUX, 67 pages, 11 planches, 1985 ............................. Fr. 


RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE 
(COTENTIN-BAIE DE SEINE, FRANCE) 
par Y. FINET, J. WUEST et K. MAREDA, 62 pages, 1991 .......................... Fr. 


GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN; 
SHELLS AND RADULAS 


by NOEINET TAWUESR and 'KMAREDA 1992 MER PEN RE ee: Fr. 
O. SCHMIDT SPONGE CATALOGUE 
par R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 pages, 49 plates, 1992 .......... Fr. 


ATLAS DE RÉPARTITION DES AMPHIBIENS 
ET REPTILES DU CANTON DE GENEVE 
par A. KELLER, V. AELLEN et V. MAHNERT, 48 pages, 1993 ...................... Fr. 


THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: 
A DOCUMENTED FAUNAL LIST 
PARMVES EINER LS 0ipases 100 RECON I EE Fr. 


NOTICE SUR LES COLLECTIONS MALACOLOGIQUES 
DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE 


par Jean-Claude CAILLIEZ, 49 pages, 1995 .................................... Fr. 
PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS 
OF ARACHNOLOGY, Geneva 1995 (ed. Volker MAHNERT), 720 pages, 1996 ....... Fr 


CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) 
(Instrumenta Biodiversitatis I) par Ivan LOBL, 192 pages, 1997.................... Fr. 
CATALOGUE SYNONYMIQUE ET GEOGRAPHIQUE DES SYRPHIDAE (DIPTERA) 
DE LA REGION AFROTROPICALE 
(Instrumenta Biodiversitatis II) par Henri G. DIRICKX, 188 pages, 1998 
A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE 
WEST PALAEARCTIC REGION 
(Instrumenta Biodiversitatis III) par Stanley BOWESTEAD, 204 pages, 1999 ............ Fr. 


THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE 
SOKOTRA ARCHIPELAGO 
(Instrumenta Biodiversitatis IV) par Beat SCHATTI & Alain DESVOIGNES, 
WF Br pases 1999 Ar En A RE LEER Fr 


35.— 
35.— 


30.— 


30.— 


30.— 


30.— 


A0 


60.— 


. 70.— 


‘A f 


Volume 107 - Number 2 - 2000 
Revue suisse de Zoologie: Instructions to Authors 


The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific 
results based on the collections of the Muséum d’histoire naturelle, Geneva. Submission of a manuscript implies 
that it has been approved by all named authors, that it reports their unpublished work and that it is not being 
considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of 
colour plates and large manuscripts. All papers are refereed by experts. 

In order to facilitate publication and avoid delays authors should follow the /nstructions to Authors and refer to 
a current number of R.S.Z. for acceptable style and format. Manuscripts not conforming with these directives are 
liable to be returned to the authors. Papers may be written in French, German, Italian and English. Authors should 
aim to communicate ideas and information clearly and concisely. Authors not writing in their native language should 
pay particular attention to the linguistic quality of the text. 

Manuscripts must be typed, or printed (high quality printing, if possible by a laser-printer), on one side only 
and double-spaced, on A4 (210 x 297 mm) or equivalent paper and all pages should be numbered. All margins 
must be at least 25 mm wide. Authors must submit one original and two copies, including tables and figures, in 
final fully corrected form, and are expected to retain another copy. 

We encourage authors to submit the text on a diskette (3,5°°, Macintosh or IBM compatible, with “Microsoft 
Word” or similar programmes). The text should be in roman (standard) type face throughout, including headings, 
except genus and species names which should be formatted in italics (or underlined with pencil); bold, small 
capitals, large capitals and other type faces should not be used. Footnotes and cross-references by page should be 
avoided. 

Papers should conform to the following general layout: 

Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces, 
name(s) in full and surname(s) of author(s), and full address(es). 

Abstract. The abstract is in English, composed of the title and a short text of up to 200 words. It should 
summarise the contents and conclusions of the paper. The abstract is followed by less than 10 key-words, 
separated by hyphens, which are suitable for indexing. 

Introduction. A short introduction to the background and the reasons for the work. 

Materials and methods. Sufficient experimental details must be given to enable other workers to repeat the 
work. The full binominal name should be given for all organisms. The International Code of Zoological Nomen- 
clature must be strictly followed. Cite the authors of species on their first mention. 

Results. These should be concise and should not include methods or discussion. Text, tables and figures should 
not duplicate the same information. New taxa must be distinguished from related taxa. The abbreviations gen. n., sp. 
n., syn. n. and comb. n. should be used to distinguish all new taxa, synonymies or combinations. Primary types must 
be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed by 
synonyms, material examined and distribution, description and comments. All material examined should be listed in 
similar, compact and easily intelligible format; the information should be in the same language as the text. Sex 
symbols should be used rather than “male” and “female”. 

Discussion. This should not be excessive and should not repeat results nor contain new information, but 
should emphasize the significance and relevance of the results reported. 

References. The Harvard System must be used for the citation of references in the text, e.g. White & Green 
(1995) or (White & Green, 1995). For references with three and more authors the form Brown et al. should be 
used. Authors’ names should not be written in capitals. The list of references must include all publications cited in 
the text but only these. References must be listed in alphabetical order of authors, and both the title and name of 
the journal must be given in full in the following style (italics can be formatted by the author): 

Penard, E. 1888. Recherches sur le Ceratium macroceros. Thése, Genéve, 43 pp. 

Penard, E. 1889. Etudes sur quelques Héliozoaires d’eau douce. Archives de Biologie 9: 1-61. 

Mertens, R. & Wermuth, H. 1960. Die Amphibien und Reptilien Europas, Kramer, Frankfurt am Main, XI + 264 pp. 
Handley, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: Wenzel R. L. & Tipton, V. J. (eds). 

Ectoparasites of Panama. Field Museum of Natural History, Chicago, XII + 861 pp. 

References should not be interspaced and, in the case of several papers by the same author, the name has to be 
repeated for each reference. 

Tables. These should be self-explanatory, with the title at the top organised to fit 122 x 180 mm. Each table 
should by typed, double spaced, on a separate page and numbered consecutively and its position indicated in the text. 

Figures. These may be line drawings or half tones and all should be numbered consecutively, and their position 
indicated in the text. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and 
lettering should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors 
should refrain from mixing the drawings and half tones. Original drawings will not be returned automatically. The 
Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents. If electronically 
scanned figures are submitted on diskettes, this should be clearly indicated on the print-out enclosed with the 
manuscript. 

Legends to figures. These should be typed in numerical order on a separate sheet. 

Proofs. Page proofs only are supplied, and authors may be charged for alterations (other than printer’s errors) if 
they are numerous. 

Offprints. The authors receive totally 25 offprints free of charge; more copies may be ordered at current prices 
when proofs are returned. 

Correspondence. All correspondence should be addressed to 


Revue suisse de Zoologie 
Muséum d'histoire naturelle 
CP 6434 
CH-1211 Geneve 6 
Switzerland. 
Phone: +41 22 418 63 33 - Fax +41 22 418 63 01 
e-mail: volker.mahnert @ mhn.ville-ge.ch 
Home page RSZ: http://www. ville-ge.ch/musinfo/mhng/page/rsz.htm 


HECKMAN 
=i 1 N C- 


SIND E pi 
Bound-To-Please" 


MAY 01 


CHESTER, INDIANA 


mr 46962 


WO ‘ine 


0 2025 


sl Fin, Pur 
7 en, 


a 


ibe aus une 


OT 
PRASRD ADS FL U ae 


terry Me 
Sos 


N 
Neo eas 
DAMEN Au à 


Sure: 
mu; 


DOTE 
SCENE 
vege 
ATAPUD AT ann Tat lio 
AN UN Rat eps Tek, 
hry RE 


ANT 
IS